DEPARTMENT OF COMMERCE AND LABOR 

BUREAU OF FISHERIES 

GEORGE M. BOWERS, CommU«lon«» 



THE MUSSEL FAUNA OF THE 
MAUMEE RIVER 



Bureau of Fisheries Document No. 757 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1912 



*» 

^^•' 



DEPARTMENT OF COMMERCE AND LABOR 

BUREAU OF FISHERIES 

GEORGE M. BOWERS, Commissioner 



THE MUSSEL FAUNA OF THE 
MAUMEE RIVER 



15 



Bureau of Fisheries Document No. 757 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1912 



A V 



X^ 



h^ > 






THE MUSSEL FAUNA OF THE MAUMEE RIVER 

By h! Walton Clark and Charles B. Wilson 



Bureau of Fisheries Document No. 757 



CON TENTS. 



Page. 

Introduction 3 

Geology and geography of Maumee and upper Wabash Basins 4 

Biology of Maumee River 8 

Mussel survey 10 

Headwaters of Maumee and Little Wabash Rivers 10 

St. Marys River and Spy Run 11 

St. Joseph River and feeder canal 18 

Maumee River 25 

Discussion of mussel species 39 

General distribution of shells in Maumee River 55 

Food of Maumee mussels 57 

Parasites of Maumee mussels 61 

General conclusions 71 

2 



T) 



THE MUSSEL FAUNA OF THE MAUMEE RIVER. 



By Charles B. Wilson and H. Walton Clakk. 



INTRODUCTION. 



This report embodies the results of an investigation of the mussel 
fauna of the Maumee River, carried on under the auspices of the 
United States Bureau of Fisheries during the summer of 1908, and 
it represents a part of the series of investigations undertaken by 
that Bureau looking toward means of continuing the suppl}^ of 
mussel shells. Upon this supply, which had begun to show signs 
of serious depletion, depends one of the important industries of the 
country, the pearl-button industry of the Mississippi Valley. 

The scope of the inquiry embraced whatever appeared to be closely 
related to mussel life, such as the history and character of the river 
and its present mussel supply as regards abundance, quality, and 
species. It was desirable also to ascertain where good breeding 
stock might be obtained, the location, character, and condition of 
mussel beds, and circumstances in which different species appeared 
to thrive. Such features of general life history as food, habits, time 
and manner of spawning, enemies, and parasites were also noted 
whenever opportunity offered. 

In the choice of the area to be studied, the features to be emphasized 
during the investigation, and similar details we have had the direc- 
tion and advice of Dr. Barton W. Evermann, Chief of the Division 
of Scientific Inquiry of the Bureau of Fisheries. His counsel has 
been of gi-eat assistance in the preparation of this report. 

In addition to the Maumee River, the upper part of the Wabash 
Basin was also investigated, in order that the two basins might be 
compared where they are close together. This latter part of the 
work also yielded many interesting results, especially in the spawn- 
ing habits of mussels and the life history of parasites. The results of 
the Wabash investigation are frequently referred to briefly where 
they carry on lines left incomplete by the Maumee studies. 

3 



4 MUSSEL FAUNA OF MAUMEE RIVER. 

The authors had as an associate in the work Mr. Donald M. Earll, 
of Washington, D. C, to whom is due much credit for the results. 

During the Maumee investigations, not only the main river, but 
also the two streams which unite to form it, the St. Joseph and St. 
Marys, were examined for the last few miles of their course, and the 
other principal affluents, the Tiffin and the Auglaize, were likewise 
investigated for a short distance above where they join the Maumee. 
Other bodies of water flowing into the Maumee or closely related to 
its basin, such as the feeder canal at Fort Wnyne and its reservoir, 
and Spy Run, a small tributary to the St. JNIarys not far above its 
mouth, were studied. Below Defiance, Ohio, the IMiami and Erie 
Canal runs parallel with the river, and was followed to avoid the 
riffles and difficulties of navigation to be encountered in the main 
stream. It otfered a favorable subject of research in itself and the 
Maumee could readily be examined from time to time, as that river 
was never any great distance away. 

During the trip to the United States fish hatchery at Put-in Bay, 
Ohio, o])portunity was taken to examine the mussel fauna of that 
region. This gave data for inference as to v\diich species had prob- 
ably ascended the Maumee from Lake Erie and which had entered 
more recently from the Wabash system. It also furnished a basis 
for comparison between the river and the lake mussels as regards 
size, etc. 

The authors are about equally responsible for the work. To Mr. 
Clark belongs the credit for the final determination of the mussel 
species, all of the observations on the food of mussels, and most of 
those upon the parasites. Dr. Wilson supervised the work and has 
furnished the geological and geographical distribution and most of 
the notes upon the various stations. 

GEOLOGY AND GEOGRAPHY OF THE MAUMEE AND UPPER WABASH 

BASINS. 

Although the Maumee River flows into Lake Erie and is therefore 
a part of the great St. Lawrence system, while the Wabash flows 
into the Ohio and thence to the Mississippi, the basins of the two 
rivers have been so intimately connected in their geological history 
and topographical features, as well as by the artificial connections 
established during the existence of the Wabash and Erie Canal, that 
they are practically one. Even at the present day the headwaters 
of the Little Wabash River approach within 3 miles of the St. Marys 
River and still nearer to small affluents of that stream, and when 
the white settlers first came to Fort Wayne they found this point 
an important portage for the native Indian tribes. There is no per- 
ceptible divide between the two basins, and a large open country 
ditch still unites them through an old flat lake plain known locally 



MUSSEL, FAUNA OF MAUMEE RIVER. 5 

as "the' prairie." A careful examination of this ditch revealed no 
mussels at all, and it is not likely they could live in the mucky 
bottom, though it is possible that glochidia might be carried through 
the stream by migrating fishes. 

During the glacial period the entire region was buried under the 
ice, and every form of molluscan life was exterminated. At the 
melting of the ice sheet great bodies of water were formed which 
could find no outlet to the eastward on account of the still unmelted 
ice, and therefore all the water was compelled to drain toward the 
south and west into the Ohio and Mississippi Valleys. The old 
channels can still be traced and are represented on the geological 
maps of the region. The geologists tell us that at first there were 
two of these lakes entirely separated from each other. The eastern- 
most one. Lake Maumee, covered the present basin of the Maumee 
Eiver and the upper part of the Wabash Basin and was 100 feet 
higher than Lake Chicago, which lay to the west. Lake Maumee 
drained through the Maumee and Wabash Valleys into the Ohio, 
while Lake Chicago found an outlet along the Des Plaines and 
Illinois River Valleys. 

The restocking of the region with mussels must have been effected 
by migration through these respective outlets. At this period sev- 
eral of the more hardy species came up through the Wabash Basin 
into Lake Maumee, as is attested by the finding of their shells among 
the fossils of the period. At the beginning, therefore, the Maumee 
Basin was restocked with Ohio Valley species of mussels, which came 
to it by way of the Wabash River." 

After the ice had retreated far enough northward to reveal the 
surface features of the region under discussion, it exposed the valley 
of an ancient preglacial river, sometimes called the Huronian River, 
and now known as the Saginaw-Grand Valley. This river valley 
runs east and west across the southern part of the State of Michigan 
and became a channel connecting the two lakes just mentioned. The 
opening of this channel gradually lowered the water in Lake 
Maumee to the level of that in Lake Chicago. A series of terminal 
moraines which had formed along the southern margin of Lake 
Maumee were enabled by this lowering of the water to assert them- 
selves and close the Wabash outlet. These moraines are known to 
geologists as the Mississinewa Moraine, to the south and west of 
which lies the Mississinewa River, the Salamonie Moraine, to the 
north and east of the Salamonie River, the Wabash Moraine to the 
north and east of the main branch of the Wabash River, the St. 
Joseph-St. Marys IMoraine to the east of those two rivers, both of 
which drained originally into the Wabash, and the Defiance Moraine 

" For an interesting discussion of tliis subject, see Simpson, American Naturalist, vol. 
XXX, p. 379-384. 



6 MUSSEL FAUNA OF MAUMEE RIVER. 

to the east of the Tiffin and Aiighiize Rivers. The former drainage 
of the St. Joseph and St. Marys Rivers through the "Wabash is in- 
dicated by the peculiar turn they make at Fort Wayne, where they 
unite to form the Maumee. This sudden change, of course, is so 
striking a feature that it has excited comment by abnost everyone 
who has considered the topography of the region. The situation is 
well expressed by Mr. Simpson who says: 

The St. Josephs, St. Marys, and Auglaize Rivers, tributaries of the Maumee, 
flow in the direction of the Wabash ; the two former join at Fort Wayne and 
flow partly bacliward as the Maumee, the whole looking like a tree with its 
branches broken down and hanging against its trunk." 

From the time of the closing up of the Wabash outlet until the 
St. Lawrence was reopened, the only outlet for Lake Maumee was 
through the Huronian channel into Lake Chicago, and thence into 
the Des Plaines and Illinois Rivers. All the mussels, therefore, 
which took part in restocking Lake INIaumee during this period 
belonged to the Illinois River fauna and entered the lake by this 
roundabout channel. The Unionida^ are essentially inhabitants of 
shallow water and would have spread along the margin of these 
glacial lakes. As the level of the lakes was gradually lowered and 
the present system of drainage was established, the mussels naturally 
followed the retreating waters, and thus finally found their way into 
the river beds where they now live. 

This account of the origin of the present mussel fauna in the 
Maumee and upper Wabash Basins is adapted from several geologi- 
cal reports of the States of Indiana and Michigan and from an 
excellent paper on the " Distribution of the Unionidse in Michigan " 
by Bryant Walker." 

From the foregoing account of the manner in which the stock- 
ing of the Maumee Basin took place, it would naturally be inferred 
that in the lists of species of mussels obtained at the various stations 
on the Maumee and Wabash Rivers by the party, there should be 
more than the usual amount of similarity between those from the 
headwaters of the two rivers, and this similarity should decrease the 
farther down the respective rivers the comparison is made. Such 
we find to be actually the case. (See p. 38.) 

But there is another factor which may at least claim a share of 
the credit for the correspondence in species. 

The Wabash and Erie Canal was in active operation between 
these two rivers for nearly half of the last century. Starting at 
Toledo, Ohio, it ran along the bank of the Maumee for the entire 
length of that river to Fort Wayne. There it crossed the narrow 
strip of country between the headwaters of the two rivers, ran along 

" American Naturalist, loc. cit. 

"Autlior's separate of a paper read before (lie Michigan Academy of Science, Mai'cli, 
1899. 



MUSSEL FAUNA OF MAUMEE RIVER. 7 

the eda^e of the old lake plain or " prairie," near the Indian portage 
and at no great distance from the Little Wabash, struck the main 
Wabash below Huntington, Ind., and then followed the latter river 
down to the Ohio. This canal was fed from the two rivers and 
opened into them at various points along its course. 

There was thus established a channel of communication which, 
during its half century of existence, must have resulted in the ex- 
change of many species of mussels, as well as other portions of the 
fauna of the two rivers. 

Such a canal furnishes an excellent breeding ground for mussels, 
as is well shown in the case of the old feeder canal at Fort Wayne 
(p. 19) and the Miami and Erie Canal still in existence between 
Defiance and Toledo (p. M). 

This artificial opportunity for interchange of species, occurring 
so recently, must have greatly supplemented the original similarity 
due to the former identity of the drainage basins. It would also 
go far toward destroying any local peculiarities that might have 
developed since the tw^o basins were separated. 

The Maumee Basin is situated in the northeastern part of Indiana 
and the northwestern part of Ohio, and covers an area of C,500 
square miles. There are no large hills in or around it, it being a flat 
glacial plain, covered with moraines and low ridges. The basin 
abounds in clay and solid subsoil, so that the river is rather muddy 
most of the time. The water is also quite hard, furnishing an abun- 
dance of lime to the mussels for tlieir shells. In the upper part of 
the river, as far down as Defiance, Ohio, there is considerable sand 
and gravel bottom, but below Defiance a good deal of the bottom 
is solid bedrock. As already stated, the Maumee is a very young 
river in point of geological time, yet, next to the Detroit River, it is 
the largest tributary of Lake Erie and is one of the largest rivers 
within the State of Ohio. The Maumee is formed by the junction of 
the St. Joseph and St. Marys Rivers, just beneath the Columbia 
Street Bridge in Fort Wayne, and flows northeast to Lake Erie at 
Toledo, Ohio. From origin to mouth is about 100 miles in a straight 
line, but as a result of the angular or crescentic form of the glacial 
moraines which separate the Maumee from the Wabash Basin, the 
river winds greatly, especially that portion of it above Defiance, and 
thus adds fully 50 miles to its length. 

As might be expected, the current is not very swift except during 
high floods, since the river falls but 164 feet in its entire course, or a 
trifle over a foot a mile, and much of this is accumulated in a series 
of riffles at Defiance and Grand Rapids. Elsewhere the river is made 
up of long, sluggish stretches, alternating with short riffles of varying 
lengths, and it is in the vicinity of the latter that most of the mussels 
are found. 



8 MUSSEL FAUNA OF MAUMEE RIVER. 

Through the removal of most of the forests which once covered the 
area about its headwaters, the current and depth of water are not so 
constant as they once were, but the river rises and falls more sud- 
denly, and is usually considerably more turbid or muddy than 
formerly. The height and activity of the river in times of flood are 
strikingly manifested by the tangled piles of drifted logs and brush 
up on the flood plain high above the normal surface of the river. 

This change in the steadiness of flow of the river, with the accom- 
paniment of a more shifting bottom, has probably had considerable 
eft'ect on the mussel life of the stream. 

f 
BIOLOGY OF THE MAUMEE RIVER. 

Through the abandonment of the Wabash and Erie Canal, which 
took place nearly 30 years ago, and the draining and gradual drying 
up of the old lake plain or prairie, the Maumee River has become 
isolated from the Wabash, so that there has been no transference of 
species for some time. 

In the course of our work our attention was devoted chiefly to the 
mussel fauna. The general features of the river and its inhab- 
itants, both plant and animal, undoubtedly have many important 
relations to the mussels, but, on account of the limited amount of 
time at our disposal and the impracticability of carrying on several 
lines of work at once, only such features were noted as appeared to 
have some evident bearing on mussel life. In the St. Marys River 
and about the region of Fort AVayne, where conditions were favor- 
able, the river was more thoroughly explored than elsewhere, the 
flora and fauna were noted, and the stomach contents and parasites 
of the mussels were examined. All along the way stomach contents 
and parasites were preserved for future study. 

A valuable addition to our knowledge of the biology of the 
Maumee Basin has been furnished by a report of the investigations 
made by Philip H. Kirsch during the summer of 1893 and published 
in the Bulletin of the United States Fish Commission (vol. xiv, 
for 1894). This report refers to an abundance of algae in various 
places, and furnishes a list of 87 species of fishes, 3 species of mussels 
in the Maumee, 4 species of crawfishes, 7 of batrachians, 6 of snakes, 
and 6 of turtles. In this report the remark is made that mussels are 
common." They are, however, only incidentally mentioned, so that 

" The mussels obtained by Kirsch during these investigations were later turned over to 
Mr. C. T. Simpson for examination. In an interesting discussion concerning the " Mis- 
sissippi Valley Unionidae found in tlie St. Lawrence and Atlantic drainage areas," in the 
American Naturalst, vol. xxx, p. 379-384, Mr. Simpson remarks of these shells : 

"This changing of characters" la number of changes have just been discussed] "has 
been well illustrated in a lot of Unionidae recently submitted for examination by Prof. 
B. W. Evermann, of the U. S. Pish Commission. * * * Unin lutcolus Lam., U. 
subrostratus Say, U. circulus Lea, l'. phaseolus Hild., U. multiplicatiis Lea, U. multira- 
diatus Lea, and Anodonta grandis Say, are so dwarfed and stunted and changed in color 
as to be scarcel.v recognizable, while the same species from tlie Wabash from which these 
have no doubt all been derived, are as vigorous and finely developed as any in the Mis 
sisslppi Valley." 



MUSSEL FAUNA OF MAUMEE EIVER. 9 

Kirsch's report and the present one may be regarded as supplemen- 
tary to each other. 

The following general observations were made concerning the 
biology of the stream : 

Among the aquatic plants, Oscillatoria was abundant in the St. 
Marys Eiver and the upper portion of the Maumee down as far as 
Kern Reservation, 7 miles below Fort Wayne. In the St. Marys 
this plant was found floating in large blackish or blue-green bunches 
on the surface of the river; " farther down much of the floating Oscil- 
latona seemed recently washed in. Filamentous algae, probably 
Cladophora^ grew on the rocks in the bottom and occasionally on 
the shells of the mussels. Wild celeiy {Vallisneria spiralis) was 
abundant on the bottom of the St. Marys at Fort Wayne, but w^as- 
not conspicuous farther down. The dense-flowered water willow 
{Dianthera am-ericaiia) fringed the edges of the river and lower 
portions of the canal all along the way and was one of the plants 
most constantly seen. During the latter part of the cruise along the 
Miami and Erie Canal this plant was covered with large tangled 
clumps of dodder, so that the prospect either way along the canal 
was lined with great masses of yellow here and there. Algaj of 
many sorts gi-ew in great abundance in the Maumee below the dam 
at Defiance. Portions of the canal from Napoleon, Ohio, to the locks 
below were covered with a dense scum of delicate little w-ater fern, 
Azolla caroliniama. Along the lower stretches of the canal Cera- 
tophyUum was abundant in stretches of quiet water. Typha latifolia, 
the narrow-leaved cat-tail, formed a dense hedgelike growth along the 
edge of the canal in places. The spatterdock, Nymphaa advena^ was 
fairly common along the lower reaches of the canal and the white 
water lily, Castalia odorata^ occasional. Duckweeds were common 
in the canal, but not noted in the river. 

As to animal life, various species of protozoa were very abundant 
in one place in the St. Marys, as will be described later on. (See 
p. 13.) They were not noted farther down the river; however, there 
was little opportunity to examine carefully for them. No plankton 
was taken, but the material examined from the stomachs of mussels 
probably gives a fair notion of the nature of the smaller free plank- 
ton elements. Sponges and flatworms were rather common on rocks 
and shells the whole length of the river. In addition to the mussels, 
%arious mollusks such as Pleurocera^ Ancylus, and Sphrenurrh were 
common. Crawfishes were abundant everywhere in the river, but were 
not often noted in the canals. xVquatic insects and larvae were not 

" Further observations show that the presence of Oscillatoria and other minute organ- 
isms is a periodic phenomenon. They accumulate during periods of low, stagnant water 
but altogether disappear during periods of high water with rapid current. 



10 MUSSEL FAUNA OF MAUMEE EIVEK, 

especially abundant, except below the clam at Defiance. According to 
residents of the region along the river, fishing was in general rather 
poor, although in a few places, as several miles above Defiance and 
below Defiance Dam, angling was enthusiastically carried on by 
men wading out in the stream in high boots and casting. In the 
autinnn of 1907 a good deal of angling was observed in the St. 
Joseph River just above its mouth, at Fort Wayne. The anglers 
there were catching a good number of the wall-eyed pike, or pike 
perch, Stisostedioit v'ltreum., which they called " pickerel.'" We did 
not see many fishes in the water during most of the cruise, although 
at Fort Wayne, Avhere we had opportunity to examine carefully, a 
number of species were observed, and all along the Maumee the 
small-mouthed black bass seemed fairly common. The darting fishes 
frequently struck the boat with resounding whacks and occasionally 
jumped over the boat or even over our heads as we sat in the boat. 
They frequently jumped against different members of the party and 
fell back into the water, and G examples, most of them quite large 
and fine, jumped into the boat and were captured. At one place, a 
little above Woodburn, Ind., we passed through a large school of 
basking gars. 

Among a(iuatic birds and waders, kingfishers {Ceryle alcyon) were 
fairly conunon, green herons {Ardea vlresceiw) and the great blue 
heron (Ardea herodias) were quite abundant, the former along shore 
or on branches of trees and the latter standing statuesquely on rocks 
in midstream and rising with a low level flight as we approached. 
We saw several small flocks of wood ducks {Aix sponsa) which seem 
to be summer residents. Sandpipers were common along shore and 
flocks of killdeer occasional on bars. On account of the northern lati- 
tude it is perhaps worthy of note that the song of the Carolina wren 
came to us frequently from the woodlands along the upper stretches 
of the river. 

No aquatic mammals were seen, although occasional shell piles gave 
evidence of the presence of a few muskrats. 

MUSSEL SURVEY. 

HEADWATERS OF MAUMEE AND LITTLE WABASH BASINS. 

The region in and around Fort Wayne is of peculiar interest, since 
it is the borderland between the two river basins, and also furnishes 
much evidence of the influence of the old Wabash and Erie Canal 
upon the mussel fauna. For this reason, considerable time was spent 
here before beginning the specific work upon the basins themselves. 
Most of the localities reported upon were worked over several times 



MUSSEL FAUNA OF MAUMEE RIVER. 11 

in order tliat there might be a reasonable certainty that all the resi- 
dent species had been found. A careful study was made of the 
species themselves and data collected as regards food, parasites, etc. 
The localities examined include the St. Marys and St. Joseph Eivers, 
Spy Run, a small tributary of St. Marys, and an old feeder canal and 
its reservoir which was formerly connected with the St. Joseph Eiver. 
The Aboit and Little Wabash Rivers, which belong to the AVabash 
system, were also examined. The stations at these localities are 
designated by lettei's, those on the main river by numbers. The rela- 
tive abundance of the different species at each station is indicated 
by giving the niunber of examples obtained, or by remarks. 

ST. MARYS RIVER AND SPY RUN. 

In the St. Marys River the shells are scattered irregularly eveiy- 
where, so that along the bank a few dead shells may be found every 
now and then, or an occasional live mussel seen in the water. There 
are a few shell beds, all of small size, however, where conditions 
appear to have been exceptionally favorable for the development of 
molluscan life, and at such places the mussels are fairly abundant. 

Station A. Above the Blujfton Road Bridge, at Fort Wayne. — The 
highest point at which the St. Marys River was examined was above 
the Blutfton Road Bridge, about 4 miles from the mouth of the river. 
At this place there were riffles and the remains of an old dam, and the 
point lies above the entrance of most of the sewage of the city. The 
water was shallow and the bottom covered with gravel, sand, or mud, 
according to the rapidity of the current. At the date examined 
(June 26) the water was quite turbid and nothing could be seen on 
the bottom. We had heard reports of a goodly number of shells 
there, but by wading about considerably we found only a few. The 
living mussels were hard to find, partly because of the stony bottom 
and partly because of their actual scarcity. A man fishing at the 
place was offering boys who were swimming near by 5 cents apiece 
for mussels, which he wished to use for bait, and it was noteworthy 
that although the boys were searching diligently the fisherman was 
not oversupplied. We found a fair number of dead shells on the 
shore, representing 18 species. 

The little Truncilla sulcata, which seemed on the whole a rather 
rare shell, and of which during the entire summer's work we found 
only 1 live example (although scattered valves were occasionally 
found along the river bank), seemed unusually common here; we 
obtained 4 entire shells and 10 valves. None of the larger species of 
mussels were present in sufficient numbers to be of commercial value. 



12 MUSSEL FAUNA OF MAUMEE EIVER. 

while all of the others were either too small or so discolored as to be 
unfit for use. 

The river was followed from this point down to Swinney Park, 
a distance of about a mile and a half, but practically no shells were 
seen, and men wading in the river fishing for crawfishes with dip 
nets knew of none. From Swinney Park down to the riffles, soon 
to be discussed, the river had been examined fairly well during the 
previous year, and no shell beds of any importance had been found. 
Occasionally dead shells had been seen scattered along shore and in 
one place a few large Lampsilis ligamentinus, which had probably 
been killed by fishermen for bait. 

Station B. Rifjles^ St. Marys River helow the Van Buren Street 
Bridge. — The most important bed found in the St. Marys River was 
at the riffles, about an eighth of a mile below the Van Buren Street 
Bridge and across from the city waterworks pumping station. 

The riffles are conveniently near the residence of Miss Elizabeth J. 
Bowman, at 719 Michael Avenue, who very kindly gave us the use of 
her house and grounds as a laboratory headquarters for the work in 
and around Fort Wayne. Being so favorably situated, this station 
was repeatedly visited and the mussels obtained were carefully exam- 
ined for food, parasites forming pearls, etc. 

As the opportunity for doing laboratory work is rather rare in 
moving field parties, and as it was supposed that this mussel bed 
would be fairly typical of others we would encounter but would not 
have opportunity to study fully, this bed was investigated and the 
observations noted in considerable detail. 

The stretch of river above the mussel bed was about 8 feet deep 
at its greatest depth and 60 or 80 feet wide, with a sandy bottom, 
and was used by the boys as a swimming-hole. A large quantity 
of sand for building purposes had been hauled out of the river at 
this place and screened some little distance from the river bank. 
Among the screenings was a fair number of mussel shells of different 
species and shells of Pleurocera and S phwriuTn. The species of mus- 
sels represented by the shells among the screenings were Qnachvda 
ruhiginosa, Q. undulata., Pleurohema clava, TJnio gihhostis, Sym- 
fhynota complanata^ Anodonta grandis, Lampsilis gracilis, L. rectus, 
L. ligamentinus, and L. luteolus. There were not many live mussels 
to be found in this stretch of the river, probably on account of dis- 
turbances caused by the hauling of sand. 

There were also few found below the riffles, as the stones were too 
thick in the river bed to furnish any foothold. The shell bed itself 
was therefore on a slope between the shallower portion of the riffle 
and the deeper sandy bottom upstream. The bottom on which the 



MUSSEL FAUNA OF MAUMEE EIVEE. 13 

mussels were located was composed of mingled sand and pebbles. 
The water over the mussel bed was shallow, averaging about 2 feet 
in depth during the period of observation, and the current was mod- 
erate, 3 or 3^ miles per hour. The water varies considerably in 
depth and turbidity at diiferent times; even a small local shower 
raises the surface perceptibly and makes the Avater very muddy, 
after wliich it slowly abates in volume and becomes clear. While 
this mussel lied Avas under observation it was occasionally rendered 
unworkable for a few days by local showers. On the other hand, the 
water was sometimes found clear enough to see the bottom with a fair 
amoimt of distinctness. 

The life conditions of the rifHes and the swimming hole just above 
it were studied in considerable detail. The river flora did not appear 
to be particularly rich. The only conspicuous alga was a species of 
OsciUatoi'ia, which was very abundant on the surface in large floating 
blue-black patches from which the filaments radiated in every direc- 
tion. Of phanerogams, the only aquatic plants growing in the river 
at this place were eelgrass or wild celery {ValUsneria spiralis) and 
the dense-flowered water willow {Dianfhera americana). 

The animal life was rich and rather varied. One of the most 
abundant animals was a V 07'ticella-\\kQ protozoan, probably Epistylis^ 
which thickly covered with a uniform hoary coating the blades of 
ValUsneria and the branches and leaves of willow which dipped into 
the water. The same or a similar protozoan grew in great abundance 
on the crawfishes of all sizes, especially thickly on the antennae, 
which were rendered quite white and plumose. Examples of the 
protozoan taken from the crawfishes showed the zooids thickly clus- 
tered at the apex of a long nonretractile dichotomously branching 
stem. On account of the nonretractility of the stem, this white coat- 
ing can not contract into a mass wdien irritated as can colonies of 
V orticella. A small colony of Vorticella w^as found clustered on an 
Atax taken out of a mussel. A species of fresh-water sponge was 
very common below the shell bed, frequently coating the inside of 
dead valves with an expanded thick feltlike covering. The brown 
hydra [Hydra fmca) was quite abundant, and numerous examples 
were seen attached to the back of shells of Ancylus. Leeches did 
not seem to be abundant ; the only one noted was a fish leech attached 
to a sunfish caught in the river. 

In addition to the mussels, discussion of which wnll be reserved 
till the end of the description of the locality, various univalves such 
as Pleurocera^ Physa, and Ancylus were common. 

The crawfishes were so abundant at this place as to deserve special 
mention. So far as examined the species was Camharus virilis 



14 MUSSEL FAUNA OF MAUMEE RIVER, 

Hagen, and the river bottom appeared to be almost overpopulated 
with them. The young were often seen crowding up close to the 
shore. As noticed above, many were thickly covered with a growth 
of protozoa, and some of the young were covered with a green algal 
growth. In the mussel bed the crawfishes were seen sunning them- 
selves on the rocks which projected out of the water, lifting them- 
selves high upon the pincers in a most peculiar fashion. Upon 
wading out to a large projecting rock situated in the river about in 
the middle of the mussel bed they were found to be clustered thickly 
on the sloping surface of the rock just at the water line, their heads 
projecting out of the water. Many were found living in dead mussel 
shells just below the bed; indeed, it was hardly possible to pick an 
old shell or tin can out of the water without obtaining a crawfish. 
Some of those thus captured were of very large size. The crawfishes 
were frequently observed devouring the flesh of dead mussels, but it 
is doubtful whether they were able to kill any, especially the larger 
ones. They may possibly be an intermediate host of some of the 
mussel parasites. The crawfishes were caught in considerable num- 
bers, either by dip nets or minnow seines, for fish bait and were used 
in the St. Joseph Eiver. On July 2 a man was observed seining 
crawfishes just below the mussel bed at the riffles. The seine was 
drawn over the bottom, which was covered with rounded pebbles 
about the size of hen's eggs or larger, and the ends of the brails 
poked about among the rocks. The crawfishes, of which it seemed 
there was one under nearly every stone, backed up into the net and 
were caught in great numbers. The fisherman wanted only the soft- 
shells and ''peelers" (crawfishes about to molt) and threw the rest 
back. A great number were molting at the time. The " peelers " 
can be easily distinguished from other crawfishes by pressing the 
sides of the carapace, which arches up free from the body. They 
also have a somewhat different color. The molted carapaces of 
crawfishes and gastroliths associated with them were quite common 
along the shore of the river. 

No crustaceans except crawfishes Avere noted in the river itself, 
though small Entomostraca were abundant at the edge of the river, 
in overflow ]">o()ls. 

Aquatic insects and larvje did not appear to be particularly abund- 
ant so far as species were concerned; a few forms, however, were 
quite common. The red "blood worm" (a Chtronom'us larva) was 
present in large numbers, and along shore, just under the water 
surface and attached to submerged objects by a short slender pedicel, 
was a great number of small gelatinous pear-shaped o])jects full of 
minute dots, probably Qgg bunches of Ghironomus. Midges were 
abundant, emerging in great numbers; on the morning of June 21 



MUSSEL FAUNA OF MAUMEE RIVER. 15 

there were many exiivite on the water surface and on objects along 
shore. Many of the insects died shortly after emerging. 

On the first visit to the swimming-hole, the afternoon of June 14, 
a large number of johnny darters {Boleosoma nigrum) and a few 
minnows were observed dead along shore. All the fishes were quite 
small, apparently the young of the year. It is very likely that some 
sudden pollution of the river about that time Avas the cause of the 
death of the fishes as well as that of a great number of large craw- 
fishes on shore and mussels in the water. Some time during the sum-] 
mer it was reported that dyestuff had been poured from the Imitting 
mills some distance above into the river and had caused a wholesale 
destruction of fishes, which died in such numbers as to render por- 
tions of the city along the river scarcely habitable. In addition to 
the dead fishes mentioned above, the following species were observed : 
Catfishes (young, probably Ameiurus natalls), mad toms {Schil- 
heodes gyrinus), golden shiners {Ahramis crysoleucas)^ carp {Gypri- 
nus carpio)^ and sunfishes {Eupomotis gibhosus). In the previous 
year a number of small black suckers {Catostomus nigricans) were 
seen at this place. 

Of reptiles, a few painted turtles {Ghrysemls marginata) were 
found. There Avere no indications of aquatic mammals, such as 
tracks or piles of shells along the shore. Late in autumn, however, 
a good many crawfishes were seen along shore with backs bitten 
open, and this may have been done by a muskrat or a mink. 

As has been said, although a few mussels could be found above 
the main bed, the greater number were to be obtained just at the 
head of the riffles in a fairly compact bed. The mussel bed itself is 
quite small, probably not more than 2,000 square feet in extent, and 
commercial operations would clean it out in a very short time. On 
several occasions the water was clear enough to see the bottom and 
the distribution of the mussels fairly well. The mussels did not 
appear to be moving. They were headed mostly downstream, the 
natural position for feeding nuissels, and were not deeply buried; 
about half of the shell projected out of the mud. The projecting 
portion was more or less stained and frequently covered Avith an algal 
growth. 

On June 20 this bed was gone over with considerable thorough- 
ness and a number of mussels collected. These were placed on shore, 
counted, and sorted over, and, after a number of representative 
shells such as Avere desired for specimens Avere picked out, the rest 
were thrown back. All the shells, especially L. rectus and L. liga- 
mentinus, were large and fine, A number of the living animals were 
measured and Aveighed, Avith the results shoAvn on the next page. 
22367"— 12 2 



16 MUSSEL FAUNA OF MAUMEE EIVER. 

Measurements and AV eights of St. Marys River Mussels. 



Serial 
No. 



Species. 



Length. 



Height. 



Thick- 
ness. 



Weight. 



Quadrula undulata 

do 

do 

do 

do 

do 

do 

Lampsilis rectus 

'.[[[do'.'.'.'.'.'.'.'.'.'.'.'.'.'.'.'.'.'. 

do 

do 

do 

do 

Lampsilis venlricosus... 

do 

do 

Lampsilis ligamentinus. 

do 

do 

do 

do 

do 

do 

do 

do 

do 

do 



do 

do 

Lampsilis luteolus. . . 

do 

do.. 

do 

do 

do 

do 

do. 

Symphynota costata. 

do 

do 

do 

do 

do 

do 



mm. 
115 
104 
102 
8(5 
80 
70 
72 
158 
159 
134 
150 
153 
141 
154 
126 
110 
83 
137 
135 
145 
135 
131 
132 
130 
126 
131 
122 
115 
97 
120 
122 
124 
127 
134 
127 
126 
118 
102 
92 
123 
95 
111 
113 
95 
111 
101 



Ounces. 
10.75 
8.25 
8.50 
4.75 
3.75 
3.50 
3.00 
13.75 
14.50 
7.75 
13.75 
12.75 
12. 50 
8. 75 
13.75 
8.25 
4.75 
16.50 
15.50 
18.25 
15.75 
14.75 
13.75 
11.75 
12.00 
12. 75 
12. 25 
10.75 
5.25 
11.75 
10.00 
9.25 
10.75 
11.00 
11.75 
10.75 
9.25 
6.50 
4.00 
6.75 
3.25 
4.75 
5.75 
3.00 
5.25 
3.50 



BeloAv the mussel bed and also in it were a large number of dead 
shells. These were covered with water and appeared to be still as 
good for commercial purposes as ever, since they retained their 
original texture and luster. Some of the shells that had been longest 
dead were stained black, perhaps from iron in the water, but this 
color was superficial and could have been easily ground off. The 
mussels appeared at the time of our visit to be dying in some num- 
bers, for a good many still retained the position they had during life 
and the flesh was still clinging to the interior of the shell. Some of 
the larger living shells were diseased and stained brown or green in 
the neighborhood of the cardinal teeth. 

The stomachs usually contained considerable mud, in which was 
scattered a few diatoms, ScenedesTnus, Phacus^ etc.; the amount of 
organic material was quite small. The results of studies of stomach 



MUSSEL/ FAUNA OF MAUMEE RIVER. 17 

contents are given in detail under the general heading at the close of 
this report. (See p. 57.) 

When examined in June these mussels were strikingly barren. 
The only gi'avid individual noted was an example of Lampsilis 
ventrivosus with the shell of a marked female contour very much 
inflated, and this was partly barren, the gills appearing shrunken. 
Subsequent investigations show^ed that the sterility of this bed was 
due partly to the season. As w^e went downstream and the season 
advanced we noticed gills of various species beginning to take on a 
dusky tinge at the margin and filling up with fertilized eggs and 
glochidia. Moreover, on a visit to this mussel bed on September 21, 
of 28 examples of L. ligamentinus procured 8 were gravid. The other 
species procured in September were : Qnadrula tuherculata^ 1 ; Q. 
ruhigiriosa, 1 ; Q. uridulata^ 3 ; Symphynota complanata^ 1 ; LampsiUs 
7'ectus, 3; L. luteohis, 2; none gravid. The percentage of breeding 
mussels was hardly as great as one might expect under favorable 
conditions. No small shells were found, the smallest being of 
medium size and apparently several years old. The conditions on 
the whole seem to indicate that this mussel bed is dying out. It 
would have been possible to secure a number, perhaps a half ton, of 
good shells here without disturbing the mussels. 

The riffles were incidentally visited in November, and although 
the water w^as quite cold the protozoa were still abundant on craw- 
fishes and dead leaves in the water, and the conditions bore con- 
siderable resemblance to those observed earlier in the year." 

None of the mussels was badly parasitized.^ Atax ypsilophorus, 
Aspidogaster conchicola^ and Cotylaspis Insignis were rather com- 
mon, but not abundant. The marginal cyst distomid was rather 
frequent, and it was here Ave obtained material with which we were 
able to associate this organism with jDcarl formation. It was most 
common in LampsiUs ligamentinus. 

From the riffles down to the mouth of the river mussels appeared to 
be quite scarce. 

" On Juno 20, 1000, as opportunity permitted, this place was revisitod. It was durin.i; 
a long period of high water. The river, though turbid, was not filthy and appeared 
reasonably clean. At the riffles it was racing at a rapid rate and the current could 
hardly be stemmed when about breast-high, but would sweep one off his feet. No Pro- 
tozoa were seen, and only a few young crawfishes, which were very active and alert. 
A few gray-back minnows were swimming about on the surface, and schools of small 
minnows darted from shore. By wading about a great deal only two live mussels (L. 
ligamentinus) were found, though there were a number of dead shells. There were no 
evidences of unhealthy conditions or overcrowding ; on the contrary, the conditions 
seemed to be those of an ideal mussel stream. This visit served to emphasize the fact 
that during periods of high water sewage is comparatively innocuous and that while the 
water of the river has great variation the river population is fairly constant ; In 
periods of low water the stream is overpopulated and, to use a chemical phrase, the in- 
habitants are almost precipitated out. 

* The parasites are more fully discussed and described at the end of the report. 



18 MUSSEL FAUNA OF MAUMEE EIVER. 

Station C. Spy Run. — Spy Rim is the name of a small shallow 
stream which enters the St. Mary's about a half mile above its mouth. 
The bottom is of fine sand and the water at the time visited was 
quite clear. There were many small fishes, chiefly the johnny darter 
{Boleosoma nigrmiv)^ a few golden shiners {Abramis crysoleucas) 
and chubs (Semotilus atromaculatus) , and a number of crawfishes, 
all free from Protozoa. No living mussels were found, but about a 
dozen dead shells, all of which, except fragments of one LampsiUs 
iris, were Anodontoides feiv/ssaeianvs snheylindraceus. 

ST. JOSEPH RIVER AND FEEDER t'AXAL. 

The St. Joseph River, more commonly known by the shortened 
local name St. Joe, is a larger and clearer stream than the St. Marys, 
and since only the very lowest portion of its course enters the city, 
it is much freer from sewage. 

Station D. St. Joseph River at Robinson Park., 6 miles north of 
Fort Wayne. — The highest point at which the St. Joseph River was 
examined was at Robinson Park, a pleasure resort about 6 miles 
above the city. This point could easily be reached by trolley. Con- 
siderable change had taken place in the river here within the last few 
years. At the time the park was established there was a dam in the 
river which had kept the feeder canal, just above, full of water. This 
had raised the water of the river and had expanded it somewhat into 
a quiet, pondlike basin. The breaking of the dam had reduced the 
water to its nornud height and emptied the canal. 

At the time of our visit the river was rather high and turbid from 
recent rains. We procured a boat and rake at the park and rowed 
up to the riffles about half a mile above the park, examining the river 
and shore. 

There was a fair number of shells scattered along here, all of 
which were large and of excellent luster. The large shells found at 
the park itself had been dead some time, and may have been killed 
by the lowering of the water when the dam broke; the LampsiUs 
hfteolus were exceptionally large and fine. Farther up the stream 
large dead shells were collected on the Inink, chiefly Quadrida undn- 
Jata. They were freshly dead and had probably been killed for bait. 
On investigating the river it was found full of snags in some places; 
in others the bottom was of hard clay, too compact for mussels to 
live in. In the beds of. gravel where mussels were likely to be found 
the bottom was full of sharp rocks wdiich in the turbid water were 
difficult to distinguish from mussels, so that collecting the latter was 
uncertain. We found very few live shells, but were told that they 
were easy to obtain when the water was low and clear. 



MUSSEL FAUNA OF MAUMEE RIVER. 19 

The manager of the boathouse at Robinson Park told ns of having 
seen a good bed of mussels here during clear low water. He also 
reported that where he used to live on the Ohio River they ate the 
" small sweet mussel," but the larger kinds were too tough for food, 
except that they could be used for flavoring soup. 

Station E. Feeder canal from Robinson Park to Fort Wayne. — In 
the days of the Wabash and Erie Canal, the dam mentioned above 
was built across the St. Joseph River just below what is now Robin- 
son Park, and a feeder canal was dug to convey the water from 
above the dam to the main canal at Fort Wayne. After the main 
canal had fallen into disuse this feeder was still used as a source of 
power for mills at the outskirts of the city, and the water was turned 
back into the St. Joseph River. 

AMien Robinson Park was established the canal formed a charm- 
ing water lane leading from the city out to it. About the summer 
of 1906 or 1907, however, the dam in the river broke and let the 
water out of the canal. 

The dry canal bed offered one of the most remarkable opportuni- 
ties possible for the study of mussel distribution within a small area. 
It was 6 miles in length and, with the exception of a few places be- 
low the general level, it dried up so quickly that the mussels are still 
left in their original positions in the mud on the bottom of the canal. 
The shells are half to two-thirds buried in the dried mud, the great 
majority of them with the posterior end directed northward — that is. 
against the current when the canal was full of water — and with the 
valves nearly closed, just as the animals died. 

This region had been visited and preliminary studies made in the 
autumn of 1907, and it presented the same features that it did early 
in the summer of 1908 — a long, dry, and cracked mud flat forming a 
vista of projecting shells, the short tracks the mussels had made 
during life being still distinct. In the canal bed it was possible to 
study all the species that had lived there, their abundance, distribu- 
tion, and the like. Many of the young shells of Lampsilis luteolus 
exhibited brilliant rays; they appear to have been exceptionally 
highly colored. 

Toward the upper end of the canal, in a place where the bottom 
was 15 feet wide, the mussels were counted for a stretch of 10 feet 
along the canal bed and the following species noted : Quadrula riib'i- 
ginosa^ 11; ^- cylindrical 1; Q. vnchdata^ 86; Anodonta grandis., 6; 
Ptychohranchus jdiaseolus., 1 ; Lampsilis ligamentinus, 5 ; L. luteolus., 
6. The width taken was the total width of the bottom of the canal 
and was considerably wider than the space occupied by the mussels. 

About a mile farther down the canal a space 10 feet square was 
measured off in the bottom of the canal and the following species 



20 MUSSEL FAUNA OF MAUMEE RIVER. 

were found: Quadi'ula riibigmom^ 6; Q. undiilata-^ 60, all rather 
small; Pleuroheina clava^ 1; Alasmidonta tnincata^ 2; Symphynota 
com'planata, 2; S. costata^ 5; Anodonta grandis, 15; Ohovaria eir- 
eulus, 4; Lampsilis ligamentinus, 5; L. luteolus, 1; L. ventricosus^ 4. 
This gave a little over 1 shell per square foot. In 1908, in a square 
meter of bottom near the Eod and Gun Club, the following sf)ecies 
were noted: QuadruJa nihiginosa^ 9; Q. undulata, 36; Symphyiwta 
complanata^ 1; Anodonta grandis^ 17; Ohovarla circulus^ 11; Lamp- 
silis iris, 2; L. ligamentimbs, 2; L. luteolus, 3, giving a total of 81 
per square meter. In addition to these shells there were many small 
Sphaniums, the ground being paved with them, 34 Campelomas, 
and 23 Pleuroce7'as. The square meter referred to above represents, 
as nearly as could be judged, an average number rather than either 
extreme. 

The general impression obtained concerning the abundance of vari- 
ous species is given in the table on page 37. 

Figure 2, plate i, represents a portion of the dried canal bed near 
its upper end. It shows admirably the mud cracks formed on drying 
and the mussel shells still protruding in situ. 

Realizing the value of an opportunity to study so extensive a tract 
with the mussels still in their original positions and in plain sight, 
several visits were made to the canal, and as a result the following 
conclusions were reached : 

1. Each of the species found was well distributed throughout the 
entire canal; that is, there w^as no marked gathering together or 
colonizing of any one species to the exclusion of others. So fully was 
this true that a careful examination of a few rods of the canal bed 
anywhere was reasonably certain to yield specimens of all the species, 
except possibly one or two that were very rare. In this connection it 
must be kept in mind that the bed of the canal was very different 
from that of a river, in that it was practically the same throughout 
its entire length. It was all shallow mud, with little sand or gravel; 
the water was of nearly uniform depth, and the current was of uni- 
form rapidity. In short, there was almost nothing except the con- 
sistency of the mud upon which a mussel could base any preference 
of locality. Slight differences in this respect caused an increase in 
the numbers of mussels in some places, but affected all the species 
equally. 

2. The number of mussels which can find accommodation under 
fairly favorable conditions upon a definite area in the canal bottom 
may be judged by the counts on measured areas given above. 

3. A canal makes an excellent breeding ground for mussels, as was 
particularly attested by the presence of a large number of younger 
examples among the adults, showing all stages of growth. The cur- 
rent in the canal appears to have been just fast enough^ food material 



U. S. B. F.— Doc. 757. 



Plate 1. 



Fig. 1. — Bed of Maumee River 
just below Columbia Street 
bridge, Fort Wayne, Ind., 
showing mussels killed by 
sewage from city gasworks. 




■^■il^C^K^^ 







liT* 



. S^ 



-*x' 



Fig. 2. — The dried bed of the 
Feeder Canal, showing mud 
cracks and mussel shells 
in situ. 







r^'^^lSy*^ 



Fig. 3.— Pile of shells, chiefly 
m u c kets . Lampsilis ligamen - 
tinus, containing 25 tons, 
gathered fromthe Maumee 
River near Fort Wayne, 
Ind. 




MUSSEL, FAUNA OF MAUMEE RIVER. 21 

constant and ample, the supply of water uniform the year round, and 
the bottom seems to have been well suited to a great variety of species. 
This gives the mussels a uniformly large size, while the shells at the 
same time are of extra quality and luster. 

4. These facts would indicate that a canal would furnish one of the 
best locations that could be obtained for artificial propagation of 
mussels. Furthermore, the water in a canal can be drawn down from 
time to time without much trouble, thus allowing examination of the 
mussels. Examples suitable for breeding or for commercial purposes 
could then be selected easily and rapidly without in any way disturb- 
ing the others. Such possibilities are at least worthy of a careful 
consideration in future attempts at the artificial propagation of 
mussels. 

The last visit to the canal was made on July 27, after a series of 
heavy rains. The canal bed was at this time under water for nearly 
its entire length, and w^as covered w^ith grass and weeds that had 
sprung up in a short time and effectually concealed the shells. This 
visit threw light on a phenomenon which had proved quite puzzling 
during previous visits. Occasionally, shells of Qvadrula undulata, 
only recently dead and with bits of flesh still clinging to them, had 
been found in dry places in the canal l)ed some distance from any 
jjool. At the time it was hard to understand how they came to be 
there, as it seemed impossible that they could have " mudded up " and 
survived the long period following the breaking of the dam. This 
visit showed that during the high water of winter or spring the 
whole canal bed had probably been submerged, and that the mussels 
had migrated from the deeper portions at that time. 

During our previous visits, also, a number of pools, comprising the 
deeper portions of the canal bed, were observed, and a few live mus- 
sels, chiefly Anodonta grandis and Qnadrula undidata, had been 
noted near the edges, but the pools were not fully investigated. 

On this last visit the pools were thoroughly investigated and were 
found to contain an abundance of moUuscan life. Fine examples of 
the folloAving species of mussels were secured: Qnadrula coccinea, 1; 
Q. ruhiginosa, many, deeply buried; Q. ci/lhidrica., 1; Q. nnduhita, 
many; AlasmMonta truncata^ 1; Symphynota costata^ several; Ano- 
donta grandis^ a few; Stt'ophitus edentalus, a few; Ptychohranchus 
phaseolus, 1; Obovai^i circulus, 1; Lampsilis iris, 1; L. rectus, 2; 
L. Ugamentinus, many, large and fine; L. luteolus, several; and L. 
ventricofius, 1, large. Among other mollusks, Ancylus was abundant 
and large, usually attached to the shells of living mussels. Algae of 
various species were common, covering the water surface. In one 
place Hydrodlctyon was exceedingly abundant. 

This investigation w^as interesting and important, as it showed that 
mussels will live for long periods in shallow pools where they once 



22 MUSSEL FAUNA OF MAUMEE EIVER. 

have obtained foothold, even though the conditions under which they 
once throve and muhiplied have ceased. While it is true that mus- 
sels die rather quickly when placed in foul water or unfavorable 
conditions, both our investigations here and our experiments in trans- 
planting elsewhere indicate that they can live for long periods where 
they probably would not naturally develop, and that they can easily 
be stored in jdooIs of limited size. 

Station F. Reservoir of the feeder canal. — This is an artificial 
pondlike area at the lower end of the canal and within the outskirts 
of the city of Fort Wayne. This reservoir, which originally con- 
nected with the canal, covers about an acre, and the water, which at 
the time of our visit was only a foot deep at most, probably never gets 
much deeper, as it can easily drain into the river below. The bottom 
of the reservoir is of a bare, yellowish clay, and there is a great 
deal of miscellaneous trash over it, indicating that it has been used 
to some extent as a sort of dumping ground. There were quite 
a number of mussels in this pond. Quadrula rubighiosa was com- 
mon in the warm shallow water near shore, and on the date of our 
first visit (June 30) was remarkably active, almost every example 
being at the end of a long, curved track. They were quite high up 
out of the mud, and several had fallen over and were lying on their 
sides. Quaditda undidata and Anodonta grandis were also common, 
and there were a few Lampsilis luteolus. Some dead L. glans and 
one dead L. venirlcosus, which quite closely resembled L. eapax^ were 
found along shore. More very young mussels were seen here than 
anywhere else. They were at the end of long, narrow tracks resem- 
bling snail tracks, in the shallow water along the north shore of the 
pond, and were nearly buried in the mud, only the posterior tip of 
the shell slightly projecting. Their presence indicates that the reser- 
voir was used quite actively as a breeding ground for the mussels. 
There were a fcAv fishes in the reservoir, chiefly carp and suckers, and 
boys were occasionally seen fishing there. Other mollusks, especially 
large Camfelorrbas.^ were abundant. 

The mussels of this place, especially the Anodontas, were more 
heavily infested with parasites than those obtained anywhere else, 
with the exception of some Lampsilis alatus taken at Grand Rapids. 
Ohio, which contained Atax in great numbers. The common mus- 
sel parasites, At(u\ Cotylaspis, and As pidog aster ^ were present in 
nearly every individual examined. 

The reservoir was again visited on July 23, and it w^as found that 
the Quadrula ruliginosa, which on the previous visit were high up 
out of the mud and actively moving about, had by this time buried 
themselves deeply. It was also noticed that at the same date, in 
the pools of the canal itself, they were likewise deeply buried. It 
is probable that their great activity during our first visit vv^as due to 



MUSSEL FAUNA OF MAUMEE RIVER. 23 

restlessness caused by the warmth of the shaHow water and that their 
action of deeply burying themselves was in anticipation of the dry- 
ing up of the reservoir. A number of the Anodontas obtained at this 
time were examined as to food and parasites, with the following 
results : 

No. 1. — Parnsites: Ala.r, 3, in gi.ll axils; Cotylastpifi infiiffnis, 1, in axil of Inner 
gills; Aspidoguf<ter conchieola, 2, in ])ericardial cavity. A small patch of Plu- 
■matella poJiimorpha on each valve. Gills empty, hut ova in the hody. Stomach 
contents ample, brownish green in mass, containing a few i^cenedesmn.s and 
numerous glohular organisms, probably Trachelemonas. 

No. 2. — Parasites: Atax, 27, in axils of gills; Cotylaspis insignis, 8, in axils 
of inner gills; Aspidogaster conchieola, 8, in pericardial cavity. Shell deeply 
stained a salmon color on the inside, with a tadpole-shaped blister on the left 
valve near the siphonal opening. Considerable PJiimatella polymorplia on the 
outside of the valve. Stomach contents fine greenish brown in mass, mostly of 
unrecognizable material, and containing several Sccncdcsmus, 1 Pcdiastrum 
pertiisum, and 1 shell of rotifer Anurwa cochlearis. 

No. 3. — Parasites: Atax, 48, between and among the gills, chiefly in the axils; 
Cotylaspis insignis, 2, axils of inner gills ; Distomuni' of Osborn about 248, on 
outside of mantle, between mantle and shell ; Aspidogaster conchieola, 8, in 
pericardial cavity; Sporocysts {Distomuni of Osborn) innumerable, distributed 
thickly throughout the mantle. Stomach contents, chietly minute green and 
black globules, probably Cryptomonas, and 1 diatom, Plcurosignm. 

No. 4. — Parasites: Atax, S large, 2 small, in gill axils. The large ones were 
full of eggs. Distommn of Osborn, 58, 30 on the right side near the umbo 
( " right shoulder " ) and 28 on the left ; Cotylaspis insignis, 9, in axils of inner 
gills; Aspidogaster conchieola, 2, in pericardial cavity. Stomach and intestine 
almost empty, but contained Trachelcnwnas, Ganwsphceria apania, and a 
spindle-shaped Euglena-Uke organism. Nacre stained a deep salmon color, with 
a good deal of Plmnutella on the outside. 

No. 5. — Parasites: Distoniunt. of Osborn, 33 on the "left shoulder" and 35 
on the right ; sporocysts of that species innumerable throughout the mantle, 
the cercarise escaping from them and present in large numbers; Atax ypsilo- 
phorus 32; Atax, small species, 18, between gills; Cotylaspis insignis 11, in axils 
of inner gills ; Aspidogaster conchieola, 5, in pericardial cavity. The intestine 
contained numerous minute dark-green globules {Trachelemonasf), which 
were moving about actively. It also contained the Euglcna-VikQ organism, and 
Anurwa cochlearis, 1 empty shell. 

No. 6. — Parasites: Distomuni of Osborn, 23 on the "right shoulder" and 
12 in the left; Atax 50, in the axils of the gills; there were many Atax eggs, 
which are minute yellow objects, on the inner side of the mantle. Two species 
of Atax present, a large and a small, both full of eggs. The eggs in the mantle 
were hatching, and the larviie were present in the different stages of develop- 
ment ; Cotylaspis insignis 4, in the axils of inner gills, and Aspidogaster 
conchieola 13, in the pericardial cavity. The shell had considerable Plumatella 
polymorplia attached; the nacre was stained a deep salmon, and there was 
a raised ridge on the right valve from the umbo downward and backward. 

No. 7. — Parasites: Atax iiitvrmedius 14, in axils of gills, and with the eggs 
scattered on the inside surface of the mantle ; Distomum of Osborn about 100, 
on the " shoulders " ; Cotylaspis insignis 9, in the axils of the inner gills ; 
Aspidogaster conchieola, 2, in pericardial cavity. The alimentary canal of this 
example was empty. Interior of the shell with a row of four salmon-colored 



24 MUSSEL FAUNA OF MAUMEE EIVEE. 

groups of knobs opposite each "shoulder" and extending from the hinge two- 
thirds of the way to the edge of the shell. Three straight ridges, also salmon- 
colored, on the left valve, radiating from the distal end of the row of knobs. 
The salmon marks in the nacre corresponded exactly with the position of the 
distomids iu the mantle. There were three bunches of distomids corresponding 
to the three ridges on the left side. 

No. 8. — Pai'asites: Distomum of Osboru, about 200 on the outside of the 
mantle at the " shoulders," but no sporocysts present. Atax intennedius 14, 
Atax aculeatus 3. and several examples of Atax ypsilophorus. The Atax con- 
tained large eggs, and all the animals were located between the gills: Cotyhifipis 
insiffiiis 2, in the axils of the inner gills; Asindogastcr conchicola 5, in the peri- 
cardial cavity. There w-as a good deal of Phnnatella itohjmorplia on the shell. 
The color of the nacre was pretty uniform, but somewhat more intense in 
the region of the unbones, where the distomids were also thickest. Alimentary 
canal full of TracheJemonas, with a few diatoms and Scenedesmus. The 
Trachelemonas were very actively swimming about, although they had been 
in the stomach of the mussels for at least 36 hours. 

No. 9. — Parasites: Ata.r 23. between the gills; the eggs abundant, and some 
young hatched; Distomum of Osborn 3, on "shoulders" as usual; Cotiihispist 
insignis 3, iu axils of inner gills; AsiridoguHtcr conchicola 7, In pericardial 
cavity. The alimentary canal contained rather numerous Scenedesmus, i 
Ffdiastrum pertusum, 4 Anurwa cochlearis, and 1 spindle-shaped Euglena- 
like object. 

In a day or two after the visit to the reservoir in which these 
mussels were obtained, it was revisited. The pond was covered with 
a green scum, but there was no opportunity to examine it more fully. 

The collections at this place were particularly interestino: because 
of the knowledae obtained concerning; stages in the life history of the 
distomid of Osborn, sporocysts of which were especially abundant 
in mussels no. 3 and 5, These matters are more fully discussed 
under our notes on parasites toward the end of this report. (See 
p. 61.) 

Station FF. Feeder canal hetween the reservoir and tlie Maumee 
River. — The feeder canal with its reservoir may be regarded as a 
sort of auxiliary channel of the St. Joseph River from Robinson Park 
down to the city. The mussels Avere, however, much more evenly dis- 
tributed in the canal bed than in the river bed. The latter was 
examined from the reservoir of the canal down to the mouth of the 
river, but only occasional mussels were obtained above Wagner Street 
Bridge. 

Station G. St. Joseph River at Wagner Street, Fort Wayne. — 
At Wagner Street there was a veiy good shell bed^ which was ex- 
amined pretty thoroughl3^ The river here is shallow, with a gravel 
bottom, and the current during ordinary stages of the water is 
about 2 or 2^ miles an hour. 

A little below this station there appeared to have been a fair bed 
of mussels; but sand was being hauled from the river in large 
quantities, and the disturbance was unfavorable to moUuscan life. 



MUSSEL FAUNA OF MAUMEE RIVER. 25 

The men hauled many shells with the sand. A fine example of Lamp- 
silis ventricdsus was observed spawning here on July 20. Just above 
where the men were hauling, Quadrula 7nibiginosa, a few Symphijnota 
costata^ some LampsiUs lufeolus, and a large number of L. Ifgamen- 
tinus were seen. Where the men were at work a number of the 
rarer shells, Pleurohemu clava and Truncilla sulcata, were obtained. 

Station H. St. Joseph Ricer jvst ahove its mouth. — One of the 
finest beds was just at the mouth of the St. Joseph River, extending 
upstream about 20 rods, and as the condition of the river was quite 
favorable this was examined several times. The bottom was of 
fine sand and the water from a few inches to about 3^ feet deep, 
and quite clear, so that the mussels on the bottom could be seen 
fairly well. There was nearly a bushel of shells on shore, which had 
been crushed by boys in search of pearls. 

The mussels had been moving about to some extent and appeared 
to be, on the whole, quite active. AMiile most of them were deeply 
buried and were headed downstream in their natural feeding position, 
a few were transverse to the current, and none showed the sharp line 
of demarcation on the shell that is usually found where the animals 
occupy the same position for a long time. Of this collection recorded 
in the table, 9 species. Q. mihiginosa, Q. pustulosa, Q. undulata, S. 
compJanata, P. phaseoh/s, L. rectus, L. Ugamentinus, L. luteohis, and 
L. ven-tricosus, represented by 208 shells out of a total of 341, were 
good button shells. This is a considerably lower percentage than a 
little farther down in the Maumee. as the proportion of marketable 
shells is considerably reduced by the abundance of the comparatively 
worthless Symphynota costata. The merchantable species were easy 
to obtain and also of large size and excellent quality. While we 
were at work on this shell bed on July 21, the clammers who had been 
working in the Maumee below entered the mouth of the St. Joseph. 
This date, therefore, marks the beginning of clamming operations on 
this river. The clammers had at the time about 800 pounds of shell 
in their boat, chiefly Lampsilis ligamentimis, which they had secured 
in the Maumee below. 

MAUMEE RIVER. 

During the autumn of 1907 preliminary investigations were made 
along the first few miles of the Maumee. Shells appeared to be 
fairly abundant, but were for the most part ignored by the local in- 
habitants. In general, no attention was paid to them except that 
they were occasionally used for fish bait, and people loiew little 
about them. A considerable pile of scattered shells was seen in a 
yard adjoining the river, and the owner of the place said he had 
gathered the mussels to feed his ducks, which were very fond of 



26 MUSSEL FAUNA OF MAUMEE RIVER. 

them. Among the shells found in the yard were some fine Lampsi- 
lis ligamentinus^ and a rather thick ventricose form of L. alatus. 
There was no clamming for button shells in the river and little or 
no pearling. 

In the summer of 1908 a prospecting trip along the first few miles 
of the Maumee discovered a clammer's camp, which will be spoken 
of more fully later on. (See p. 27.) The camp was visited several 
times during preliminary field work. 

The boat was launched July 29 just below the Columbia Street 
Bridge in Fort Wayne, which marks the head of the Maumee, and 
August 9 we reached Miami, Ohio, 7 miles above Toledo, where the 
river enters Maumee Bay. The party proceeded downstream by 
easy stages, examining the river's bottom for mussels wherever there 
were indications of their presence, stopping overnight at one of the 
towns or villages along the river or at a convenient farmhouse. De- 
fiance and Grand Eapids, Ohio, were the only places where any stay 
was made. At Defiance the Tiffin and Auglaize Rivers enter the 
Maumee, each of them nearly as large as that stream itself, and 
three days were spent examining them. At Grand Rapids an extra 
day was devoted to the long riffles below the dam. The old Miami 
and Erie Canal is still in operation from Defiance to Toledo. This 
afforded a way of getting past the dams and riffles and gave promise 
of being itself rich in mussels. As it kept so close to the river that 
the latter could be visited easily at any time, the party chose the 
canal for the remainder of the trip. The following is a summary of 
the stations with the species found at each and their relative 
abundance. 

As previously stated, the stations were numbered consecutively, 
and the numbers appearing with the species represent the specimens 
of each that were obtained. 

Station 1. Maumee River at its souree. — This includes the first 
2 or 3 miles of the river and the clammer's camp. 

At the veiy head of the river, just below the Columbia Street 
Bridge, an immense bed of dead shells was found, forming banks and 
bars. The entire bottom of the river and the south bank for some 
distance out of the water Avere literally paved with shells, chiefly 
muckets (LampsHh ligamentinus). 

Some idea of the number of these shells can be obtained from 
figure 1, plate i, in which every one of the objects upon the bank is a 
dead shell. In all probability these mussels had been killed by the 
refuse from the gas works which are situated on the bank of the St. 
Marys a few rods above its junction with the St. Joseph. Spots of 
tar were found on dead mussels some distance below this point. The 
water was covered with an oily scum in places and a tarry odor was 
perceptible for several miles down the river. 



MUSSEL FAUNA OF MAUMEE RIVER, 27 

The bottom of the river at its head is Devonian limestone, and the 
banks are of white chiy; the average depth of the channel at the time 
examined was about 3 feet and the current about a mile an hour. 

A mile and a half down the river was the clammer's camp. This 
was the only one on the river and had been established but a short 
time. The owner, Mr. H. S. Birge, had formerly lived on the 
Wabash, where shelling operations are active, and had done con- 
siderable clamming there. He had come to Fort Wayne to Avork on 
the construction of a canal, and finding this work slack at times^ had 
naturally turned to the river and begun investigating its clamming 
possibilities. He reported clamming " fairly good but not as good as 
one might expect from the looks of the river." As the bottom was 
rough and stony with occasional snags and the water fairly shallow, 
he procured shells in the same fashion that the}' are obtained in parts 
of the lower Wabash — by scooping them up with a coal fork, the 
operator standing Avaist deep in water. In three weeks' time, with 
the help of his brother and two young boys, the clammers had 
gathered 25 tons of hrst-quality shellSj mostly *' muckets " (Lampsilis 
Ugamentinus) , with a sprinkling of good "long Johns" or "black 
sand shells" {Lampsilis rectus)^ a few " pocketbooks " (Z. ventri- 
cosus). etc. The shells were remarkably uniform in size and quality. 
There were few culls or worthless shells — less than a bushel. They 
consisted chiefly of Lampsilis alatus^ a few Qiiadnda tuherculata^ 
and an occasional Z^nio gihhosus. 

P'igure 3, plate i, shows a pile of shells, with the cleaning table in 
the background, and gives a fair idea of their size. In the matter of 
pearls and baroques this lot of shells gave very meager returns ; there 
were no baroques worth saving, and only two small and comparatively 
worthless pearls in the whole lot. Mr. Birge was impressed with 
the size and fine quality of the muckets of this region. 

Station 2. Five miles below Fort Wayne. — A fair bed of mussels 
was found here, with many dead shells along the bank. The water 
was shallow with almost no current, the bottom muddy with an 
abundance of alga:". 

Station 3. Kern Reserration. — There was here a large bar project- 
ing out of Avater and covered Avith dead shells Avhich had evidently 
been washed out during riA^er floods. The riAcr Avas free from sew- 
age and the Avater Avas quite clear, about 3 feet at the deepest, and 
Avith very little current. The bottom Avas of rounded graA'el well 
coA^ered in places with algpe, and many good-sized sponges. There 
were also large numbers of flatAvorms on the stones and many 
Ancylus, while crawfishes were numerous and hostile. 

The mussels were well bedded and quite abundant, but not A^ery 
large. A sample collection Avas preserA-ed, after which the others 
were throAvn back. This was the first station at Avhich Q. pustulosa 



28 MUSSEL FAUNA OF MAUMEE EIVER. 

was obtained in abundance, although a few fair-sized examples were 
found in the feeder canal in 1007. Mr. Kern, whose farm extends 
along the river here, says that the mussels appear to be dying off 
more rapidly than formerly, due, he thinks, to increased pollution of 
the river by sewage. 

Station If. Near New Haven^ Ind. — Just above the New Haven 
Bridge is a long stretch of the river with clear shallow water and 
gravelly bottom. Here a fairly thick bed of mussels was found. 
The crawfishes were especially numerous and pugnacious, and many 
of them Avere seen eating small mussels. It could not be ascertained 
whether they had killed the mussels or were merely feeding on some 
that had died from other causes. All the shells obtained were rather 
small. The Q. undidata were peculiar in that the ridges were con- 
siderably broken up into isolated portions somewhat resembling 
tubercles. 

Symphynota com-phmata, which was one of the characteristic shells 
of the feeder canal and of which wo obtained examples in the mouth 
of the St. Joseph, may have been introduced into the Maumee River 
through the agency of the Wabash and the Erie Canal. At the town 
of New Haven we were told of a remarkable shell that some boy had 
found along the river several years before, and which had been pre- 
served as a curiosity. An examination proved it to be a character- 
istic specimen of this species. This was the farthest cloAvn the 
Maumee it was seen or heard of. 

Station. 5. Below Netv Haven., Ind. — This station was near a sand 
bar just below a long riffle. The water was from a few inches to 3 
feet deep, and the bottom sandy, with many pebbles. There was an 
immense number of crawfishes here, one of which was eating a dead 
Q. undidata. Only a few live mussels were found. Among the dead 
shells were one Alasmidonta truncata^ one Strophitus edentulv^, 
several Ohovaria circulus, and several Lampsilis rectus. 

Station 6. One m/ile heloio station 5. — A small pile of shells was 
found on the river bank opposite a house owned by a man named 
Nieter. Some boys had been searching for pearls and had thrown 
the emptied shells together. Upon examination we found only three 
species. 

Station 7. Fovr mUes above Woodbrirn, hid. — After leaving station 
G no indications of shell beds were noted until we reached the vicinity 
of Woodburn, Ind. Four miles above that town is a large gravel 
bar on the north bank of the river, overflowed at high water and 
covered with dead shells. Several live mussels were found in a pool 
on the bar and one or two w^ere out of water and still alive, showing 
that they had come there during the recent high water. The list 
includes both living and dead shells. 



MUSSEL FAUNA OF MAUMEE RIVER. 29 

Station 8. Woodburx, Ind. — Near ^Vooclbul•n is a long series of 
riffles knoAvn as Bull Rapids, where shells appear to be abundant, but 
would be difficult to obtain, except during very low water, on ac- 
count of the rocky bottom and the swiftness of the current. There 
was a long bar here covered with dead shells, but although we waded 
about everywhere over the coarse sand and rocks of the bottom, cover- 
ing a large area, we found only a few live mussels. The party 
stopped for the night with a Mr. Armbruster, just below this station, 
and he reported finding in his cornfield on the south bank of the 
river a large pile of mussel shells some distance from the water, and 
made up of the same kinds as those found in the river. The presence 
of bones and stone implements among the shells left no doubt that 
this was an Indian kitchen midden, and that the aborigines had used 
these mussels for food. 

Station 9. Mawniee Center Bridge. — Just above Maumee Center 
Bridge some men were bus}^ hauling gravel out of the river bed. 
There was a fair number of shells in the gravel, though by no means 
enough to indicate a shell bed, and the men were of the opinion that 
there were not many in the river. The water was turbid at the place, 
the bottom of gravel, and the depth 3^ to 4 feet. The species noted 
were found in the sand along the shore. 

Station 10. One mile below Antwerp^ Ohio. — The next place at 
which the river was examined was about a mile from Antwerp, Ohio. 
Just below the town the river has a width of 250 or 275 feet, with 
loAv banks, and there is a long series of riffles. The bottom is covered 
with coarse gravel and loose rocks. There were many dead shells on 
bars alongshore, while the live mussels were rather scattered and 
deeply buried among the rocks and pebbles. 

Station 11. Five miles below Antwerp^ Ohio. — This station was 
along the bed of the river opposite the farm of a man named Curtis. 
A long sand bar with a number of dead shells was seen. On the bar 
were noted a few Qiiadriila rubiginosa, several Q. eylindriea^ a few 
Q. iindulata.1 a large number of Pleurohema clava^ a few TJnio gib- 
bosKS, a few Obovaria circidus, several Lampsilis iris, a few L. rectus, 
and a large number of L. ligamentinus, which was the most common 
shell. Just above the bar the Avater was deep and sluggish and be- 
low it was shallower and swifter. In the deeper water the shells 
were widely scattered and cleeph^ buried and consequently difficult 
to find. In the shallow water on the slope of the bar they were also 
well scattered, but had been activel}' crawling about and were easily 
found at the end of the tracks the}' had made. 

Station 12. Ten miles ahove Defiance, Ohio. — The water was quite 
deep here and the river banks steep; the place was used as a swim- 
ming hole. The bottom was rather solid, composed of fine mud and 
sand and covered with a growth of algae, among which was a large 



30 MUSSEL FAUNA OF MAUMEE RIVER. 

handsome species of Chara. Xo shells were seen on shore, and after 
examining a large area of the bottom only 3 live mussels could be 
found : 1 Unio gibbosus, 1 Lampsilis alatus, and 1 L. luteolus. 

Station 13. Tiffin River ^ just abore Defimice^ Ohio. — Tiffin Eiver is 
a large stream entering the Maumee about a mile and a half above 
Defiance, Ohio. We had heard that there Avere mussels in the stream, 
and Kirsch'* has reported them as abundant in the lake at the head of 
the river, although he says nothing of the river itself. The back- 
water of the Maumee, caused by the dam below Defiance, extends for 
about 2 miles up this river, so that it was a very sluggish stream 
with almost no current. A light-green thin scum, resembling 
Euglena, flowed slowly down, streaking the central portion of the 
river. 

The banks and bottom were yellow clay, the water turbid, and so 
far as we could observe aquatic vegetation was lacking. "We as- 
cended the river about 2 miles. No shells were seen on shore and 
considerable dredging brought up only 3 live mussels, 1 Lampsilis 
alatus, 1 L. ligame?itinus, and 1 L. ventricosus. The river was full 
of brush and snags which frequently entangled the dredge. It is 
quite likely that mussels are more abundant higher up beyond the 
dead water. 

Defiance, Ohio, Avas reached August 2. Here the Maumee and 
Cincinnati Canal goes southward through the State. At the time of 
our visit this canal was nearly empty, as the locks were being re- 
paired. This canal was followed on foot for some distance into the 
country, but no traces of mussels of any kind could be seen along 
the shore or in the mud in the bottom. 

Station U. The Avglaize Rircr, 2 miles abo-ve Defame, Ohio.— 
The Auglaize River is fully as large as the Maumee, and joins the 
latter at Defiance, the old Indian fort for which the city is named 
having stood upon the point of land between the two rivers. The 
water is considerably clearer than that of the Maumee, and the dead 
water backs up the river for some distance. Just beyond the dead 
water is a series of short riffles. Here the water was quite shallow 
and the bottom composed of a slippery horizontal shale which had 
been used to some extent in the manufacture of cement. The river 
is further broken up by a number of small islets and pools. Many 
large dead shells were found on the bars. The mussels could obtain 
no foothold in the bottom of the main portion of the river, since it 
was of bare shale, but were found in the sand and gravel bars along 
the banks. Here they were congregated into dense beds. Nearly the 
whole surface of the bars was covered with dead and living shells, 
even among the roots of the water willows and other aquatic plants. 
The water was clear and the current moderate, making the collection 



o Bulk-tin U. S. Bureau of Fisheries, loc. cit. 



MUSSEL FAUNA OF MAUMEE KIVER. 31 

of specimens easy. Shells of all the species found were large, some 
of them exceptionally so, being the largest of their kind we had yet 
seen. This was notably true of Sy7npkynota costata and Alasmidonta 
tvuncata. The shells were also of the finest quality and best luster, 
making them very desirable material for commercial purposes. 

The L. vent7'icosus were buried rather deeply and were compar- 
atively hard to dig out. Quadnda pustulosa^ which had been 
dwarfed thus far in the Maumee, was here of good size. It will be 
noted that several species were found here which had not been en- 
countered before, viz, Quadrula lachrymosa^ Plagiola donaciformis 
and elegans^ and Ohliquana re-fiexa. (See fig. 5, pi. ii.) 

Station 15. The Auglaize River^ 4 i^Hes above- Defiance. — The suc- 
cess at station 14 determined us to try the river again 2 miles farther 
upstream. The bottom was much the same as at the previous station, 
except even more slippery and with fewer islets; and the only place 
the mussels could obtain any foothold was in sheltered coves along- 
shore, now on one side and now on the other, according to the cur- 
rent and swing of the river. The coves differed somewhat as to 
bottom, some being sandy or gravelly and others muddy. The shells 
found here included more of the smaller species than at the preceding 
station, although there were also some very large ones. On one of 
the fine gravelly bars a number of mussels, chiefly of the smaller 
species, were very actively crawling about in the shallow water; 
a recent shower had probably moved them out of their places and 
they were readjusting themselves. Here was found the only living- 
example of Trtmcilla mdcata seen during the whole summer's work. 
In the protected coves with muddy bottom Atwdonta grandis was 
the most common shell. Lampsilis ligamentinus was just beginning 
to show an approach to the gravid condition ; the lower edges of the 
outer gills were becoming faintly distended and margined with 
black. PtychohrancJius phaseolus was considerably further ad- 
vanced, the gills being thrown into wavy folds. There were many 
small schools of minnows here, which kept nibbling at our feet and 
ankles when we waded into the stream and which clustered eagerly 
about bits of bread which we threw into the water. 

Among mussel parasites the marginal-cyst distomid was quite 
common, and there were a few Atax and Cotylaspis present. The 
Anodontas were heavily infested with Atax and the distomid of 
Osborn. 

Station 16. Maumee River ^ 4 miles below Defiance. — Four milas 
below Defiance a State dam, 7 feet high, crosses the river. This 
dam, known also as the Independence Dam, was built by the State 
of Ohio to make the river a feeder to the Miami and Erie Canal, 
which enters the river at this point. In the palmy days of the 
22367°— 12 3 



32 MUSSEL FAUNA OF MAUMEE BIVEE. 

canal the town of Independence was located here at the dam, but 
when the railroad superseded the canal the town moved elsewhere. 
The water above the dam backs for 7 miles up the Maumee and about 
2 miles up the Tiffin and Auglaize Rivers and forms a charming 
lakelike body of water below Defiance. This water is navigable 
for small boats and launches, of which there were a considerable 
number. The river is some 600 feet wide at the dam, and at the 
time of our visit a thin sheet of water was pouring over. We had 
been informed while at Defiance that there were many fine mussels 
below the dam. (See fig. 4, pi. ii.) 

Examination of the place proved the accuracy of our informant 
and showed that this locality was the richest collecting ground and 
supported a greater variety and abundance of life than any other 
stretch of the river. The river bed below the dam is a broad valley 
with a limestone bottom, the river being broken up into a great num- 
ber of pools and channels dotted with little islands overgrown with 
willows. The water, sparkling and fresh after its race over the 
dam, seemed to make conditions exceptionallj'' favorable for all sorts 
of aquatic life. The shores of the pools were edged with water wil- 
lows, and the stones on the bottom were covered with a rich growth 
of algse of various species and hues, some growing in long ropy 
masses; there were especially large clumps of Cladophora^ lending 
a green color to the bottom, and numerous larvae of caddis and stone- 
flies gluing the pebbles together. Many crawfishes were seen, and 
Bryozoa were common on the rocks. There were many anglers in 
high boots wading about in the racing currents. Flocks of black tern 
were flying overhead, the first we had seen. 

Fringing the edges of the pools and islands were many rather small 
shells, chiefly Plar/iola elegans and young Qvadrula lachrymosa that 
had apparently been killed by fishermen for bait. By feeling about 
in the sand in the shallow water shells could be found in great num- 
bers, and they were of fine quality and very large size. The L. ven- 
tncosus were the largest we had seen. Everything considered, these 
are the most important mussel beds in the river, and this is one of the 
places worthy of future study. 

It was at this ph\ce we found a couple of pearls, which, on account 
of their minute size, were of no market value. They were, however, 
of considerable interest, as, en account of their position, they served 
to intensify the suspicion already entertained that pearl formation 
may be excited by the presence of the marginal cyst. We are of the 
opinion that this is an exceptionally favorable location for the carry- 
ing on of investigations along this line. 

At the Defiance Dam we entered by means of a lock into the first 
stretch of the canal, which runs from Defiance to Grand Rapids, Ohio. 



MUSSEL FAUNA OF MAUMEE RIVER. 33 

The canal has a moderate current and is a picturesque waterway, with 
a winding towpath on one side and a country road on the other. 

Station- 17. Maumee Ricer at FloHda Biidge^ Ohio. — This is -1 
miles below the dam, and the river had very much the same character 
as at the previous station, except that there appeared to be more water 
than at the very foot of the dam. There was not a great deal of cur- 
rent, and the water was clearer than any we had yet seen ; the mussels 
could be clearly distinguished in 18 inches of water. The bottom 
was still of flat rock, so that it was only in favorable places that the 
mussels could anchor. They were fairly abundant, but considerably 
smaller than at the dam. 

Station 18. Miami and Erie Canal., near Texas., Ohio. — Xear a small 
village named Texas considerable dredging w^as done in the canal. 
The uniformity of bottom and absence of rocks or snags made the 
dredging very easy. The bottom was of soft black mud and the water 
about 5 feet deep with a moderate current. AVe found here a scat- 
tered bed of Quadrula undidata; every dredge haul brought up a few 
examples. These were large and r.ather plump, with a black epider- 
mis, and approached more nearly to the form of Q. 'plicata than the 
very compressed brownish examples we had been finding hitherto. 
These shells bore colonies of PluniateJla polymorpha. Below Texas 
there are two tedious locks in the canal, after which we entered the 
broad expanse of the Maumee above Grand Kapids Dam. At the 
last lock we saw valves of Q. lachri/mosa. 

Station 19. Maumee River above the Grand Rapids Dmn. — Here, 
as at Defiance, the dam backs up the water for several miles, and at 
this place the river is widened to 2,000 feet or more. Mussels were 
reported plentiful in this dead water and a few hauls wdth the dredge 
verified the correctness of the report. The water here is 18 feet in 
depth and the bottom is covered with mud in which are patches of 
sand and rocks, and it is very good dredging, being free from snags. 
Only three species were obtained, but they were all large, fine shells. 

Station 20. Maumee River below the Grand Rapids Dam. — This 
dam, like the one at Defiance, was built by the State of Ohio as a 
feeder for the IMiami and Erie Canal. It is 5 feet in height and a 
little over 1,800 feet in length and interrupted at the center by an 
island 350 feet in width, known as Purdys Island. At the time of 
our visit (Aug. 17) no water w^as running over the dam, all the sur- 
plus escaping by the canal on the north bank of the river and a mill 
race on the south bank. The river bed below the dam is broken uj) 
into numerous channels separated by islands covered with water wil- 
lows and other plants much as it was below Independence Dam at 
Defiance. Here, however, the bottom is a fine-grained sandstone 
which has been largely quarried for building purposes, with the 



34 MUSSEL FAUNA OF MAUMEE EIVER. 

result that the channels are broken by long deep ruts and nnmerons 
pools. The water was very clear and in the shallower portions of the 
pools the shells could be seen distinctly. In one of the pools espe- 
cially the mussels, crowded together and interspersed with numerous 
crawfishes, showed up so clearly that they made a recognizable im- 
pression on a photographic plate through about a foot of water. 

The mussels here evidently have a hard time finding enough soil 
to hold them in place, and many appear to be Avashed out and down 
the river during high water. Several were seen lying on their sides 
on the bare ledges. (See fig. 4, pi. ii.) 

There was some pearl excitement at Grand Kapids, but it appeared 
to be confined chiefly to boys. The mussels can be very easily ob- 
tained, and shells, cut open by the pearlers, Avere strewed in large 
numbers over the rocks about the edges of the pools. Several of 
the boys that we saw had vials of small irregular slugs or baroques 
to sell, but they contained nothing of any value. The inhabitants of 
the place told us that button men had been there prospecting for 
shells and had offered $40 per ton for a species which we judged from 
their description to be the black sand shell, Lmnpsilis rectus. The 
shells of that particular species were so uncommon, however, that 
nobody thought it worth while to hunt for them. 

There was some fishing in the pools below the dam, and mussels, 
especially small Quadrula pustulosa^ were used for bait. 

Station '21. Mumii and Eine Canal at Grand Rapuls^ Ohio. — At 
the end of the dam opposite Grand Rapids we entered into the second 
stretch of canal, which leads from Grand Rapids to Toledo, Ohio. 
At the very head of the canal we dredged and examined the shores 
for signs of mussels. "We obtained only one Anodonta grandis on 
the dredge, and this was heavily infested with the sporocysts of the 
distomid of Osborn. A few dead Pl-agiola elegans and Ohliquaria 
refexa were found on shore, probably left there by a muskrat. The 
canal is a pleasant stretch of water Avithout any obstructions, run- 
ning parallel with the riA^er. (See fig. 0, pi. ii.) 

Station 22. Miami and Erie Canal^ 4 nviles helow Grand Rapids. — 
About 4 miles down the canal, where a long fringe of upland forest 
borders the canal opposite the towpath, we came upon half a bushel 
of shells at the end of a burrow occupied by a mink or muskrat. 
The shells Avere all small. The P. donaciform'/s Avere of fine color 
and unusually large size. The finding of this pile of shells was a 
fortunate circumstance, as it contained a good number of shells be- 
longing to species Avhich we could not find in any quantity and 
which are not readily obtained by the dredge. Indeed, considerable 
dredging in the canal opposite the shell pile yielded only Q. undulata 
and Q. lachrymosa. 



U. S. B. F.— Doc. 757. 



Plate ||. 



1 




?^^' 




i 


._. .^i^mr^^m^'^^ ^" . 



Fig. 4. — The Maumee River 
below the State Dam, De- 
fiance, Ohio, the richest 
collecting ground in the 
river. 



Fig. 5.— The Auglaize River, 
2 miles above Defiance, 
Ohio, showing the small 
Islets and pools containing 
dense beds of mussels. 





Fig. 6.— A stretch of the 
Miami and Erie Canal, an 
excellent breeding ground 
for mussels, and an ideal 
place forartificial propaga- 
tion. 



MUSSEL FAUNA OF MAUMEE EIVER. 35 

As a usual thing, the muskrat appears to feed on other material, 
chiefly vegetables, during the summer, beginning its nuissel diet more 
extensively in the autumn, and the first mussels killed are usually 
small. 

From this station on, piles of shells along the shore indicated mol- 
luscan life, which would otherwise have escaped notice, and also the 
presence of muskrat s. 

Station 23. Miami and Erie Canal^ 8 miles helow Grand Rapids^ 
Ohio. — Here, opposite a farm belonging to Mr. Neifer, a few hauls 
were made with the dredge, and a single Quadimla pustidosa and 15 
to 20 Q. nndidata were obtained. 

Station 21^. Maumee River, Jf miles above Waterville, Ohio. — The 
river here is mucli wider than at Grand Kapids; there are broad 
riffles and the bottom is mostly of flat limestone. There are only a 
few patches of sand in the bottom among the numerous islands, 
which are covered with willows, water willows, etc. Shells were 
fairly plentiful where they could obtain a foothold. There are many 
dead shells on the islands and a few small U. gihhosus and P. elegans 
on shore freshly killed, probably by a muskrat. 

Station 25. Miami and Erie Canal, opposite station 2^. — Two 
dredge hauls were made in the canal opposite the last station on the 
river, but only a single example of Anodonta grandis was obtained. 

Station 26. Miaini and Erie Canal, 1 mile east of Waterville, 
Ohio. — Just below the bridge which crosses both the canal and the 
river, 1 mile east of Waterville, Ohio, two dredge hauls were made. 
The canal here was about 6 feet deep and the bottom was of a rather 
soft black mud. The Q. uiidulata were rather inflated, with black 
epidermis and well covered with the bryozoan Plumatella, like those 
taken near Texas, Ohio. 

Station 27. 3Iiami and Erie Canal at Westcott, Ohio. — At West- 
cott, Ohio, a small village above Miami, a few dredge hauls were 
made in the canal, but only one Quadrula lachrymosa and a few 
Q. undulata were obtained. 

At Miami, which is only 7 miles from Toledo, the Maumee was 
wide, with but few islands, and deep enough to be navigable for 
launches, of which there were many in evidence. There were large 
patches of Nelumho lutea scattered about in the river. The shore 
was examined for a considerable distance but no shells were found. 
During low water the riffles are said to extend from Grand Rapids 
to within a quarter of a mile of Miami ; the shells, therefore, would 
probably be about the same for the entire distance. 

Station 28. Miami and Erie Canal helow Miami, Ohio. — About a 
mile below Miami a temporary dam had been thrown across the canal 
to allow the making of certain repairs in the locks below. This dam 



36 MUSSEL FAUNA OF MAUMEE EIVER, 

stopped all the water except a small overflow stream which trickled 
along the very deepest part of the canal. At one place, where a 
small stream came rushing over a miniature cataract, a dense school 
of young fish, which appeared to be the skipjack, PomoJohus chryso- 
chloris^ were crowded together making frantic efforts to leap up the 
fall. 

In the shallow water of the canal many mussels were found alive, 
while along the exposed portions of the canal bottom an opportunity 
was presented for studying mussel distribution similar to that in the 
feeder canal at Fort Wayne. 

By far the larger part of the bed of the canal was open to observa- 
tion, the central stream being very narrow. At one place the canal 
when full had overflowed at one side, forming a large lagoon or 
backwater, in wdiich the w^ater had been from G inches to a foot in 
depth. The bottom of this lagoon was nearly covered with dead 
shells. 

It was noticeable that QuadTula undulata and Q, pvstulosa with- 
stand a scarcity of water better than most of the other species. There 
were more of these two species alive in the water than all the others 
together, and in several places on the exposed sides of the canal 
examples of one or the other of these two species w^ere found with 
the flesh still adhering to the shells, showing that these mussels had 
withstood drjnng up longer than associated species. The little 
Anodonfa imhecillis was found for the first time in considerable 
numbers at this place, both in the deeper and shallower parts of the 
canal. The conditions of the canal also gave an excellent oppor- 
tunity to get very young shells of some of the smaller species, such 
as PlagioJa elegans and P. donadfornds. 

In connection with the last station on the river may be considered 
shells obtained on the beach of Put-in Bay Island in Lake Erie. 
These represent species of the lake into which the Maumee flows and 
from which that stream may have received some members of its 
mussel fauna. No live mussels could be obtained at Put-in Bay 
except a few occasionally washed up by waves. About a mile below 
the bathing beach a number of dead shells were obtained, represent- 
ing the species given in the table. 

In this list the words " common " and " few " refer only to the 
relative number of shells found. Considered as a whole, shells at 
Put-in Bay were rather rare, and all seen were much dwarfed. 



MUSSEL FAUNA OF MAUMEE EIVER. 



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88 



Mussel fauna op maumee eivee. 



W 3 




MUSSEL FAUNA OF MAUMEE RIVER. S^ 

DISCUSSION OF MUSSEL SPECIES. 

1. Qitadnda fuherculata (Rafinc^^qne). Purple warty-back. This 
is not a very common species of mussel in the Maumee or its tribu- 
taries, only a few occasional examples having been taken. It was 
most abundant in the upper portions of the river, near Fort Wayne, 
and down as far as Maumee Center Bridge. No living examples 
were obtained below Defiance, Ohio, and only a very few dead shells 
were picked up below here, the last one being obtained at Waterville, 
Ohio. All specimens seen were kept, and only about 52 were obtained 
in the Maumee system altogether. The range of this species is given 
by Simpson" as "Mississippi drainage generally; southern Michigan; 
San Saba County, central Texas." While it is in the main a Missis- 
sippi form, it is one of those species which occur occasionally within 
the St. Lawrence Basin, and although not usually a lake-dwelling 
species a few old valves w^ere found along the shore of Put-in Bay, 
The Maumee examples show very little variation. Although most of 
them were rather small, some of the shells seen at the clammer's 
camp near Fort Wayne w^ere quite large. The few mussels of this 
species obtained alive usually contained small baroques in the dorsal 
region. 

This mussel is of no value for buttons on account of its color. In 
addition to being purple, the luster is usually dull. The epidermis, 
which is commonlv brown, is somewhat tinged with green in examples 
from Maumee Center Bridge. In some places the distomid of Kelly, 
associated with dorsal baroques, is a common parasite. The examples 
found in the Maumee River, however, are comparatively free froifi 
this parasite. 

2. Quadrula coccinea (Conrad). This species is not as common in 
the Maumee Basin as Q. tvhercvlata^ only about 31 specimens having 
been obtained. It was more abundant in the upper course of the river, 
but dwindled out and finally disappeared; the greatest number (10) 
was obtained at Kern's Reservation, not far below Fort Wayne, and 
the last one was taken at Florida, Ohio, in the ritlles below the bridge. 
It is a species common to both the Mississippi and the St. Lawrence 
drainage. The larger number of examples seen were white nacred, 
though a few were rosy. 

The larger white-nacred specimens would make excellent button 
shells, classing with the pig-toes, though considerably thinner. 

3. Quadrula ruhiginosa (Lea). Quadrula rubiginosa is one of the 
most widely distributed of our fresh-water shells and occurs in a 
great variety of situations; it is found in lakes as well as in rivers. 
It is fairly common all along the Maumee and its tributaries (except 
the Tiffin, which was not thoroughly examined and in which few 

" Proceedings U. S. National Museum, vol. xxii^ p. 795. 



40 MUSSEL FAUNA OF MAUMEE RIVEB. 

slie41s of any species were found). It was very abundant in the feeder 
canal and in its reservoir. A number of shells of this species were 
also found on the shore of Put-in Bay Island, Lake Erie, 

Although during the summer's collection we obtained material 
exhibiting considerable variation, all examples found within the 
Maumee Basin were fairly constant in form, though some were short 
and humjDed. This species very closely resembles Quadrula cocci nea, 
the only apparent ditierence being in the prominence of the ridge on 
the posterior umbonal slope; hence some specimens are difficult to 
classify. 

Young shells of this species exhibit faint rays which disappear 
with age. The specimens obtained in the reservoir dilfered from 
most of the others in having a very satiny epidermis. The greater 
number of the shells have a white nacre, although in some cases it is 
yellowish or rosy. None of our examples are as rosy, however, as 
Q. cocc'inea frequently becomes. The flesh of this species varies from 
salmon or orange to deep red, the richness of its color probably vary- 
ing with the nearness to the breeding season and the abundance of 
ova present; these latter are of a bright carmine color. Some speci- 
mens were found becoming gravid and one nearly ripe in the feeder 
canal July 27. On account of its commonness not all of the shells 
seen were collected; however, we obtained 112 examples from the 
Maumee Basin. 

What seemed a peculiarity in the action of this mussel was the 
habit of burying itself in the early summer. On a visit to the 
reservoir of the feeder canal on June 30, the mussels were all pro- 
jecting far out of the mud and actively moving about, while on 
July 23, at the same place, they were buried quite deep, and hard to 
find. They were also deeply buried in the feeder canal pools about 
the same date. 

A probable explanation of this peculiar action is that the reservoir 
was quite shallow and the water became warm. The water becomes 
low during drouths, and the mussels may have gone down into the 
mud in anticipation of the pool going dry. 

This species is sometimes more or less affected with Atax^ though 
not badly. Those found in the reservoir were more affected than 
others. 

Although its small size is something of a disadvantage, this species 
would make a fair button shell. It has good luster, and its compara- 
tive flatness would permit its being worked up economically. 

Though there are a number of river shells superior to it, it is one 
of the best if not the best button species that would thrive well in 
ponds. A considerable number were weighed and measured, and 
the largest, obtained at the Defiance Dam, was 88 millimeters high 
by 306 millimeters long, and the dry shell weighed 101 grams. 



MUSSEL FAUN^A OF MAUMEE RIVER. 41 

4. Quadnda piiHtulo.sa (Lea). Pimple-back: '.varty-back. Qiiad- 
Tula pustulosa is another of the widely distributed mussels common 
to both the Mississippi and St. Lawrence drainage systems; but, 
although thus widely distributed, it is somewhat choice of its habitat. 
It shows a distinct preference for more or less sandy or gravelly 
bars with a live current. It does not usually form beds, but is gen- 
erally more or less mixed in with other species. It was found 
throughout the length of the river, but most abundantly in the cen- 
tral portion. A few examples, some of them of good size, were 
found at Fort Wayne, especially in the feeder canal. The greatest 
number taken at one place was at Kern's Reservation, where -tC were 
obtained in a shallow bottom of sandy gravel. All these examples, 
and, indeed, nearly all found in the Manmee until we reached the 
Auglaize River, were small, dwarfed shells; those found in the 
Auglaize River and near the dam at Grand Rapids were of consider- 
ably larger size. A few of the shells were almost devoid of pustula- 
tions, and one small smooth shell, much produced forward and 
rounded, bore considerable resemblance to Ohovana retusa. The 
two largest examples obtained were 85 millimeters long and weighed 
(^dry shell) 145 grams. 

This is one of the species commonly used in button making, and 
where it reaches large size is fairly well adapted to that purpose. 
In the Maumee, on account of both their small size and their fewness, 
they are not an important item. Most of the examples secured were 
retained and numbered in all 156 shells. 

5. Quadrulci lacliryniom (Lea). Stranger; monkey-face. This 
species was found only in the lower courses of the river; none were 
encountered above the Auglaize. In that stream three examples, the 
first seen, were secured. From this point they were occasionally met 
with all the way down the river, and in the canal. Thirteen speci- 
mens were taken below the Defiance or Independence Dam, and below 
the dam at Grand Rapids, Ohio, 24 were obtained. This is the 
greatest number for any locality. The largest one seen was obtained 
here; its length was 115 millimeters and its weight (dry shell) 221 
grams. All examples seen were retained, the total number being 46. 

This shell is used in button making, and ranks in this respect along 
with the three-ridges. The irregular sulcation of the valves, how- 
ever, puts it to some disadvantages in cutting, so that while it is an 
interesting and well-marked shell, it does not possess enough favor- 
able qualities from the commercial point of view to make it desirable 
for propagation. 

6. Quadrula cylindrica (Say). Rabbit's-foot. This has been re- 
garded as a species of the Mississippi drainage, belonging -principally 
to the Ohio, Cumberland, and Tennessee River systems. Only a few 
examples were found and these were in the feeder canal and upper 



42 MUSSEL FAUNA OF MAUMEE RIVER. 

parts of the Maumee. It is therefore probable that they represent 
the result of migrations by way of the Wabash and Erie Canal. One 
example was found in the St. Marys River and four in the Maumee 
in the vicinity of Antwerp, Ohio ; a few dead valves were occasionall}" 
seen, but not more than two or three at the most. None were found 
below the point just mentioned. In the feeder canal, where condi- 
tions were particularly favorable for a study of the fauna, four exam- 
ples were found, and two fine valves, not mates, were obtained in the 
mouth of the St. Joseph. The shells found in the feeder canal were 
of an elegant compressed form with numerous triangular tubercles 
on the anterior portion, and represent the subspecies striglUatu^ 
B. H. Wright, which is considerably more attractive than the type 
form. They are rather dark in color, with green triangles, as is 
usual, and fine capillary rays. One example contained two Atax. 
This attractive species is not well suited to button making on account 
of its form and the fact that the nacre is frequently diseased ; hence 
it is practically unknown to the trade. 

7. Quadi'ula undulata (Barne^:) . Three-ridge, blue-point. With the 
exception of Lampsilis ligamentinii^, Quadrida undulata is the most 
abundant species of mussels in the Maumee Basin. All the undulate 
mussels found were referred to this species except those at Put-in Bay, 
Lake Erie. Notwithstanding the statement made in nearly all the 
literature at hand that '"the two species" are perfectly distinct and 
there is no need of confusing them," many examples were found that 
were hard to place. Different authors do not agree perfectly as to 
the differences between these species and, generally speaking, the 
characteristics given are rather vaguely expressed, a fault not due to 
the carelessness of describers but to the nature of the case. All de- 
scriptions seem to agree in stating that the umbones are more inflated 
in plicaia, but we find this feature as varial)le as any other. In the 
large collection of shells of the Bureau of Fisheries belonging to 
these two types we have many intermediate forms and several series 
exhibiting almost perfect transition from one species to the other. 

This species, as we have it, is exceedingly variable, so much so that 
in looking over the entire summer's collection it was possible to give 
the locality of a number of shells simply from their appearance. 

(J. imdulata was found to be fairly connnon along the river where 
there were any shells at all, and on account of its abundance only a 
small proportion of the shells seen were retained. Fourteen were 
obtained at the riffles in the St. Marys, where it was fairly common. 
In the St. Joseph Eiver at Robinson Park and in the feeder canal it 
was abundant and attained large size; 10 examples were obtained 
from the St. Joseph and 23 were taken of the large number seen in 
the feeder canal. It also throve in the reservoir of the feeder canal. 



" Q. undulata and Q. plicata. 



MUSSEL FAUNA OF MAUMEE EIVER. 43 

Very few were found in the upper Maumee. None were noted in the 
shell pile collected by the clammers near Fort Wayne and only 10 
were obtained in the river above Defiance, Ohio. In the Auglaize 
River the species is quite abundant and 52 specimens were secured. 
It was fairly common in the Maumee below Defiance. Ten examples 
were secured at the Defiance Dam, 6 in the canal at Texas, and 16 
near the dam at Grand Rapids. It was the most abundant shell in 
the canal below Grand Rapids. 

From Defiance down the river and in the canal these shells were 
l)Iacker and considerably more inflated than those above; the speci- 
mens found in the feeder canal at Fort Wayne were usually quite 
compressed and brownish in color, while those below Defiance were 
full and black, suggesting Q, plicata. These latter shells were usually 
overgrown with patches of the bryozoan Pliitnatella polymorpha. 

This is one of the staple species used in blitton making, the nacre 
having a good luster and the shell being thick and heavy. The small 
end is beautifully iridescent and frequently of an attractive blue 
color. 

This species, along with its relatives, is in good repute among pearl 
hunters. We found a few minute pearls and small rough baroques, 
but no more than in other shells. It is fairly free from parasites; 
a very few Atax were found. The nacre is frequently stained brown. 
As the species is hardy and thrives in a variety of conditions, espe- 
cially in canals, it might be a good sort of propagate where better 
species would not thrive. The total number of examples obtained in 
the river was about 180. 

In Lake Erie, at Put-in Bay, we found a dwarf species of shell, 
Q. plicata hippopma (Lea), resembling this. 

8. Pleurobeina clava (Lamarck). Simpson" regards this as a 
shell of the Mississippi drainage and gives its range as " Ohio, Cum- 
berland, and Tennessee River systems, reported from Iowa City, 
Iowa, St. Peters River, Minn., and from Nebraska." Call reports 
it from the St. Marys and from another stream in Indiana (St. 
Joseph) of St. Lawrence drainage. In the Maumee Basin it was 
nowhere abundant, but fairly well distributed along the upper course 
of the river. One was found above Blulfton Road Bridge, and in 
the St. Joseph River two dead shells were found, one quite large. 
Two shells were found in the Maumee near Fort Wayne, while at 
Maumee Center Bridge we obtained 12 moderate-sized specimens. 
At Antwerp two small shells were seen, and dead shells were still 
fairly common several miles below that place. None were seen as 
far down as Defiance, nor any below there, nor were any foimd in 
Lake Erie at Put-in Bay. The whole number of specimens obtained 

» Synopsis of the Naiades, Proceedings U. S. National Museum, vol xxii^ p. 746. 



44 MUSSEL FAUNA OF MAUMEE EIVEE. 

was 20. The Maumee specimens are brightly polished and can be 
distinguished by this feature from the rest of the summer's collection. 
This is an exceedingly handsome shell, but of no commercial im- 
portance, as both its small size and peculiar form are disadvan- 
tageous. The nacre, though usually white, is occasionally of a 
beautiful warm pink. 

9. Unio gibbostis {Burnes). Spike. Unio gihhosu^ was f Airly com- 
mon the whole length of the river. It is one of the species of wide 
distribution and considerable variability. Our specimens do not 
exhibit much variation, however. Most were of the solid, short- 
ened, somewhat humped type. The long examples, which have pro- 
cured for the species the name " spike," sometimes closely re- 
semble Lampsilis rectus, from which they can be distinguished by 
the sculpture of the umbones and by the shape of the lateral teeth, 
while the short humped form is frequently almost indistinguishable 
from Ptychobranchus phaseolus; about the only way to tell them 
apart is by observing the color and texture of the epidermis, that 
of U. gihhosus usually being darker, and by the shape of the body 
cavity. One shell was found in the St. ]\Iarys, two in the St. Joseph 
near its mouth, and three in the Maumee near Fort Wayne. It was 
most abundant in the central portion of the river from Maumee 
Center Bridge to the bridge at Florida, Ohio, and was common in 
the Auglaize River. Lower down the Maumee it gradually disap- 
peared, the last few shells having been seen at Grand Rapids, Ohio, 
below the dam. In all, 63 shells were obtained in the Maumee Basin. 
It appears to be one of the common shells in Lake Erie, 16 having 
been found on the shore of Put-in Bay Island. Both purple and 
wdiite nacred forms were present. The greater number, about two- 
thirds, had white nacre, one had rosy nacre, one had purple nacre, 
white at the edges, and the others were intermediate between w^hite 
and purple. Young examples showed delicate rays. 

This species is of no commercial value, as even the white forms 
have the nacre quite dull. In some localities the distomid of Kelly, 
associated with dorsal baroques, is a common parasite, but they are 
not abundant in the Maumee. 

10. Alasmidonta truncata (B. H. Wright). This is not an espe- 
cially common shell in the Maumee Basin. According to Simpson" 
it is a Mississippi Valley species which has migrated into the upper 
parts of the St. Law^rence Basin. It is rather variable in form and 
is sometimes difficult to distinguish from StTophitus edentulus. It 
was found in the feeder canal and St. Joseph River at Fort Wayne. 
Occasional examples were encountered along the course of the river, 
as 2 at Kern Reservation, 1 at Maumee Center Bridge, 1 at Ant- 
werp, Ohio. Three large ones were obtained in the Auglaize River 

" Synopsis of the Naiades, Proceedings U. S. National Museum, vol. xxii, p. 671. 



MUSSEL, FAUNA OF MAUMEE KIVER. 45 

ind three in the Maiimee below the Independence Dam. The hist 
example was found below the dam at Grand Rapids, Ohio. In all, 
15 specimens were taken. 

This shell is handsome and odd, brightly splashed with blue or 
green and greatly inflated. I^ is of no commercial importance on 
account of its thinness. The nacre is usually whitish, though fre- 
quently with a bluish tinge and occasionally a beautiful pink or 
rosy. 

11. Symphynota complanata (Barnes). Heel-splitter; hackle- 
back. This species is common about Fort Wayne, being especially 
abundant in the feeder canal, where it was one of the most con- 
spicuous members of the mussel fauna. Of the great number seen 
in the feeder canal 18 fine examples were taken; 2 were secured in 
the reservoir of the feeder canal and 5 in the St. Joseph River near 
its mouth. 

This species is evidently an overflow from the upper Wabash 
drainage. Only one specimen was obtained in the Maumee River 
and that was at the very head near Fort Wayne; one which had been 
found at New Haven, about 7 miles below Fort Wayne, had been 
kept and was shown as a curiosity. 

This shell was formerly not much used in button making, as it is 
usually rather thin for that purpose. Its broad flat surface and fair 
luster, however, are greatly in its favor, and more buttons can be cut 
out of one of its valves than from any other species except the large 
Quadrula heros. It appears to breed abundantly and bring forth 
large numbers of young. Gills of examples obtained in the Little 
Wabash River near Fort Wayne, September 26, 1906, were full to 
bursting with glochidia, which are brownish and give the whole mass 
a distinct brown color. 

The species appears to have bred successfully in the feeder canal, 
as there were a great many you.ng shells there. It would probably 
not thrive well in ponds, though a healthy example was seen in the 
reservoir of the feeder canal. It is not found in lakes, but is fre- 
quent in small streams with rather muddy bottoms, and is somewhat 
similar to Anodonta in habits. It is frequently the host of Atax^ 
and is one of the species especially susceptible to infection by the 
parasite Aspidogaster conchicola^ which inhabits the pericardial 
cavity, without, however, doing much apparent harm. 

As this species lives where there is rather little current and thrives 
on a muddy bottom, it would probably be a valuable subject for culti- 
vation. Its abundance in the canal suggests that it would respond 
well to artificial conditions. 

12. Symphynota costata (Rafinesque). Fluted-shell ; squaw-foot. 
This mussel was fairly common in the INIaumee River throughout 
most of its length. One example was found in the St. Marys River 



46 MUSSEL FAUNA OF MAUMEE EIVEE. 

above the Bliiffton Road Bridge, and 11 were obtained at the riffles. 
In the feeder canal 2 examples were secured. It was probably not 
uncommon there, but on account of its general abundance no more 
were taken. From 1 to 3 were found at most of the stations on 
the Maumee, so that it would be unnecessary and tedious to give its 
occurrence in detail. It was common in the Auglaize, where 13 ex- 
amples were secured, and abundant in the Maumee below the dam 
at Grand Rapids, Ohio, where we obtained 23. One was found in 
the canal just below Grand Rapids. Some of .the examples are 
beautifully rayed. The nacre is usually of a peculiar and pleasing 
yellowish shade. On account of the usual thinness of the shell, how- 
ever, the species is commercially unimportant, few or none being used 
in the button business. Altogether we obtained in the Maumee 
Basin 125 shells belonging to this species. 

13. Symphynota- com.'pressa (Lea). In the Maumee Basin we 
found only one example of this widely distributed species. This was 
found in the St. Joseph River near its mouth. The species is rather 
small and thin, and of no commercial importance. 

14. Anodontoides ferussacianiid sicbclyindraceiis (Lea). This 
subspecies was found in only one locality in the Maumee drainage, 
and there it was the only species found except one fragment of Lamp- 
silis iris. This was in a small stream, Spy Run, which enters the 
St. Marys at Fort Wayne. In this stream seven examples were 
found, all small dead shells. This species is of no commercial impor- 
tance on account of the thinness of the shell. 

15. Arwdonta grandis (Say). This species delights in a muddy 
bottom and a slow current. It was rather common in such places as 
the feeder canal and its reservoir, and a few large examples (dead 
shells) were found along the St. Joseph River near Robinson Park. 
It was found only occasionally^ in the rivers; 5 were obtained at the 
riffles of the St. Marys River; 10 were taken of many found in the 
feeder canal ; 12 were collected in the reservoir of the feeder canal. 
We found 8 in the St. Joseph River near its mouth. In the Maumee 
it occurred rather sparingly, 1 example having been obtained at the 
Mau.mee Center Bridge, 10 in the Auglaize River, 1 in the Maumee 
below the dam at Defiance, 5 below the dam at Grand Rapids, and 2 
in the canal below Waterville. A number were seen below the dam in 
the canal at Maumee. Only a few of the examples seen were col- 
lected ; the total number taken was 51. 

The specimens from the feeder canal were unusually thick, appear- 
ing as if a second nacreous layer had been deposited. 

Of all the species of mussel encountered, Anodonta, was the richest 
in parasites, harboring the greatest number of species, and (except 
in the case of some Lampsilis alatus obtained at Grand Rapids which 



MUSSEL. FAUNA OF MAUMEE EIVER. 47 

contained great numbers of Atax) the largest number of individual 
parasites. 

Most of the shells obtained were more or less stained red interiorly. 
This stain, caused by a parasite, the distomid of Osborn, varied 
greatly in extent, beginning in the umbonal cavity and extending 
from that region in two diverging lines, one anteriorly and the other 
posteriorly, ventrad. The V-shaped area thus formed gradually fills 
in with red until the whole surface is involved. This process has 
been quite fully described by H. L. Osborn « and by H. M. Kelly.^ 
The minute distomids, showing pink against the mantle, were present 
in numbers varying from a few to several hundred. 

Occasionally mussels of this species were found in which the dis- 
tomids mentioned above had not yet escaped from the sporocysts. 
The shells of such mussels frequently had the nacre raised in sharp 
pearly projections, and in some cases the sporocysts seemed to have 
been overgrown with the nacre of the shell. 

In addition to the distomid of Osborn, A. grandis also usually has 
a number of Aspklogaster conchicola in the pericardial cavity, several 
individuals of Cotylaspis insignis in the axils of the inner gills, and 
one or more species of Atax among the gills; even leeches are fre- 
quently found within the shells. The leeches are no doubt parasitic, 
as is evidenced by the fact that where a pile of freshly cleaned shells 
killed by muskrats were found under water on another occasion (at 
Lake Maxinkuckee) the leeches almost covered the insides of the 
valves. 

On account of the thinness of its shell, Anodonta grandis^ and 
indeed all the species of this genus, are of no value commercially. 

16. Anodonta imbeeiUis (Say). This species has not heretofore 
been reported from the Maumee Basin. It was only infrequently 
found; one specimen was obtained on the shore of the St. Marys 
River, at Fort A¥ayne. After this none were seen until at the very 
end of the investigation, in the canal at Maumee, Ohio, below the 
dam; at this place 38 examples were taken. Their abundance here 
indicates that they were probably abundant also throughout the 
length of the canal, but do not " bite " at the dredge hooks. As none 
were found in the bottom of the feeder canal, it is probable that they 
do not occur there. This is a very pretty species, but it is of no com- 
mercial value. 

17. Strophifus edentuhts (Say). Squaw-foot. This species is not 
common in the Maumee Basin. Examples were occasionally found 
in the St. Marys River at Fort Wayne, in the feeder canal, and in the 
St. Joseph and upper Maumee Rivers. One was obtained at Grand 

"Zoological Bulletin, vol. i, no. 6, 189S. 

"Bulletin Illinois State Laboratory Natural History, vol. v, p. 400, 1899. 

22367°— 12 4 



48 MUSSEL. FAUNA OP MAUMEE EIVER. 

Eapids, below the dam. None were seen in the Auglaize River, nor 
in the Maumee at Defiance. One shell was found on the shore of 
Put-in Bay, Lake Erie. In all, only 8 examples were obtained in 
the Maumee Basin. It is of no commercial importance. 

18. Ptychohranchus phaseolus (Hildreth). This species was not 
abundant anywhere in the Maumee Basin, but was scattered all 
along the length of the river. In the autumn of 1907 a fair number 
were obtained in the feeder canal. We found 16 good specimens in 
the St. Joseph River at Fort Wayne, 2 in the Maumee near its head, 
and 2 or 3 at occasional stations along the river. It was fairly com- 
mon in the river below Antwerp, Ohio, where 10 shells were obtained 
at one station. Nine examples were secured in the Auglaize River, 
2 in the Maumee below the dam near Defiance, and 3 at the bridge 
at Florida, Ohio, and 2, the last seen, were taken below the dam at 
Grand Rapids, Ohio. The examples obtained in the Auglaize August 
8 were beginning to become gravid. In all, 53 specimens were secured 
from the Maumee Basin. Shells of this species were fairly common 
on the shore of Put-in Bay, where it was small but beautiful in color, 
luster, and form. The outer gills of gravid examples are remarkable 
for their folded and plaited appearance; the gills look like nests from 
which insect larva^, like " Avigglers," are ready to emerge. The nacre 
of some of the young shells is of a beautiful faint pink, but that of 
all the large old shells was white. 

This shell has not been used to any extent, if at all, probably on 
account of the scarcity of supply, and it is not known to the trade. 
Its flatness, nearly uniform thickness, and excellent luster would 
make it worthy of experimentation in the button business. It would 
probably rank with the " butterfly," Plagiola secvris, in that respect 
The presence of a number of young shells in the feeder canal indi- 
cates that it might be easy to propagate in artificial channels. 

19. Ohliquaria re-flexa (Rafinesque). Warty-back. This species 
was first encountered in the Auglaize River, where 1 example was 
secured. Below the Defiance Dam 3 were obtained; 1 was found at 
Florida Bridge, and 3 at Grand Rapids below the dam. In the 
stretch of canal below Grand Rapids it was apparently abundant. 
One was picked up on the shore at the vei-y entrance of the canal 
and 22 in a pile left by a mink or muskrat along the side of the canal. 
A few were found in the canal below the dam at Maumee. This 
species, being quite thick and of good luster, is used to some extent 
commercially, but on account of its small size a shell yields only a 
few buttons. 

20. Plagiola donaciformis (Lea). The first example of this species 
was obtained in the Auglaize River. No others were seen until we 
reached Grand Rapids, Ohio, where 1 was found in the Maumee 
below the dam. In the stretch of canal near Grand Rapids it ap- 



MUSSEL FAUNA OF MAUMEE EIVER. 49 

peared to be fairly common; 10 shells were taken from a pile left 
by some predaceous animal, and of a large number seen in the canal 
below the dam at Maiimee 14 were secured, some of them quite 
young and small. 

This is one of the most beautiful of our fresh-water shells, and 
the markings are very attractive. It is too small to be of any im- 
portance commercially. 

21. Plagiola elegans (LiQn). Deer-toe. No examples of this species 
were seen until the Auglaize River was reached, where we found 10. 
It was fairly common in the river and canal below this point. Living 
mussels and shells of some recently killed were very common in the 
Maumee below the dam near Defiance. 

Of many examples seen we obtained 17 shells here. Below the 
dam at Grand Rapids, Ohio, we secured G shells, and at the pile of 
shells near the canal bank, Avhich has been frequently referred to, 
we obtained 49 shells of this s])ecies. Some were seen along the 
shore of the Maumee 4 miles above Waterville, and it was common 
in the canal below the dam at Miami. 

This is one of the prettiest of our fresh-water bivalves, but it is 
too small to be used much for buttons, though the largest examples arc 
sometimes utilized. By clammers it is called the "deer-toe." It is 
fairly constant in form, but varies considerably in color and mark- 
ings, the ground color being sometimes yellowish and sometimes 
green. The rays are often broken up into V-shaped blotches. The 
total number of examples collected was 97. 

22. Plagiola securis (Lea). Butterfly. Of this species only 1 ex- 
ample, a large coarse specimen closely resembling some of the short- 
ened and flattened forms of Lam^psilh U game tit hius, was secured. 
This is a species belonging to the Mississippi drainage, and it is one 
of the most valuable shells for buttons. The specimen was obtained 
at the clammer's camp near Fort Wayne and was from the upper 
part of the Maumee. 

23. Ohovaria circulus (Lea). There appears to be considerable 
confusion among authors regarding the three so-called species Oho- 
taiia circuluSy 0. letu^ and O. leihii, and by some authorities the 
two last named are regarded as subspecies of the first. We obtained 
the small inflated 0. leibil on the shore of Put-in Bay Island. In 
the Maumee we obtained flatter, larger shells, Avhich are similar to 
those in the collection of the Bureau from the Ohio and Tennessee 
Rivers. This species [circuliis) occurs occasionally throughout the 
length of the Maumee. Three examples were found in the St. Marys 
River at Fort Wayne. It was fairly common in the feeder canal, 
where 16 specimens were secured, and in the St. Joseph River near 
its mouth, where we obtained 10. Six examples were obtained at 
Ivern Reservation, five at Maumee Center Bridge, and 14 in the 



50 MUSSEL FAUNA OF MAUMEE EIVER. 

Auglaize River, in addition to occasional shells picked up here and 
there. The last one seen was at Grand Rapids below the dam. The 
whole number obtained in the Maumee Basin was 60. This species 
is frequently infested with the distomid of Kelly, and contains small 
dorsal baroques. Diseased shells with one or both valves roughened 
and yellow on the inside were quite common. The shell is usually 
too small to be of much use commercially, but it has a beautiful 
luster, Avhich continues in dead shells when exposed to the weather 
longer than in any other species observed. The nacre is usually 
white, though in some instances it is pink or rosy. 

24. LampsiUs gracilh (Barnes). Lamps His gracilis is fairly com- 
mon and well distributed in the Maumee Basin, but is nowhere par- 
ticularly abundant. Seven examples were obtained at the riffles in 
the St. Marys River and three in the St. Joseph near its mouth. In 
the upper part of the Maumee River it is rather scarce; only 1 was 
taken near Fort "Wayne, 1 at Maumee Center Bridge, and 1 below 
Antwerp, Ohio. In the Auglaize River it was fairly common; 13 
were obtained there. In the Maumee below the dam near Defiance 
it was rather abundant, 10 being found at that place. It was also 
abundant at Grand Rapids beloAv the dam ; here 10 were secured, and 
2 in the canal below Grand Rapids. It was quite common in the 
canal below the dam at Miami, Ohio, though none were collected 
there. 

In all, 63 examples were secured, and 10 additional dwarfed speci- 
mens on the shore of Put-in Bay Island, where it was quite common. 
This is one of the most iridescent shells we have, the posterior por- 
tion shading into a fine rainbow blue and purple. It is of no com- 
mercial value, however, on account of the thinness of the shell, which 
usually cracks badly in drying. 

25. Lampsih's alatus (Say). Pancake. This species is of occa- 
sional occurrence in the Maumee Basin, but not abundant. Along 
the upper parts of the basin they were rather rare. Three were 
obtained in the St. Joseph River at Fort Wayne, and 1, small, in the 
INIaumee, not far below its source. In 1907 several examples were 
obtained in the Maumee near Fort Wayne. A few scattered speci- 
mens were found below Antwerp, Ohio, and 1 in the Tiffin River 
a half mile above its mouth. Three were taken in the Auglaize and 
11 large ones below the dam near Defiance, where it was beginning 
to become common. Three were found at Florida Bridge. Below 
the dam at Grand Rapids, Ohio, it was one of the most common 
shells and great numbers that had been opened by pearlers were 
seen on the rocks. We secured 23 examples here. A few were 
found in the second stretch of canal and in the Maumee above Water- 
ville, Ohio. It was common in the canal below the dam at ]\Iiami, 
Ohio. In all, 56 specimens were taken in the Maumee Basin. At 



MUSSEL FAUNA OP MAUMEE RIVER. 51 

Put-in Bay Island, Lake Erie, 10 small dwarfed examples were 
found ; it appears to be common there. 

The JMaumee Basin examples, especially those obtained in the 
upper course of the river, differ from the usual type of the upper 
Mississippi Eiver specimens in being considerably thicker, shorter, 
and more inflated. Aspidogaster conchicola is a fairly common 
parasite in the pericardial cavity, and the examples seen at Grand 
Rapids were infested with Atax to a remarkable degree, some in- 
dividuals harboring nearly a hundred of the mites apiece. 

There is a great deal of variation in the color of the nacre of this 
species, from pink to rich coppery red. Some of the shells were 
beautifully iridescent. On account of its color and the thinness of 
the shell it is of no use for buttons. 

In the upper Mississippi, where this species thrives in great num- 
bers, it forms the bulk of the cull shells left on shore, and on beds 
devoted to commercial purposes would probably be regarded as a 
nuisance. Eichly colored baroques, usually of small size, occasionally 
occur in this species. 

26. LampsUis glaths (Lea). This small, inconspicuous species is 
very likely to be overlooked in collecting, and is probably more com- 
mon and more generally distributed than reports would indicate. It 
inhabits both ponds and rivers. In the Maumee Basin only 6 
examples were secured — 4 in the feeder canal, 1 in the reservoir, 
and 1 in the St. Joseph River near its mouth. All were dead shells 
picked up on shore. 

27. Lojnps'dis iris (Lea). This attractive little shell does not 
appear to be common in the Maumee Basin. Two examples were 
secured in the St. Marys River above Bluffton Bridge, 4 in the St. 
Joseph near the mouth, 2 in the feeder canal, and 1 at Maumee 
Center Bridge, making 7 in all. It is probably more common 
than collections would indicate, as it is frequentl}^ found in abundance 
late in the fall after muskrats have begun collecting, where it has 
been difficult to find in numbers before this. It seems to be one of 
the favorite articles of diet with the muskrat. 

28. Lampsills rectm (Lamarck). While not particularly abun- 
dant, this is a fairly common and well-distributed shell in the Mau- 
mee Basin. Its distribution is so general that it is unnecessary to 
give details. It was quite common in the riffles in the St. Marys, and 
present in the feeder canal, St. Joseph River, along the Maumee, and 
in the Auglaize. Nearly every example seen was kept. In all, 63 
were taken in the Maumee Basin and 3 in Lake Erie at Put-in Bay. 
Only a few young were seen, and a few gravid ones were obtained. 
The young are more or less distinctly rayed ; in the adult the epider- 
mis is about uniformly black, but the rays are represented by fine 



52 MUSSEL FAUNA OP MAUMEE RIVER. 

plaitings. The female shells are more or less inflated and pro- 
duced postero-ventrally, so that the ventral outline is somewhat 
arched. 

Generally speaking, the color of the nacre is variable in this 
species; in some localities many of the shells are pink or purple. The 
great majority of the shells in the Maumee Basin are white, though 
the cardinal teeth may be pinkish, pinkish-purple, or purple. Coty- 
laspis is a frequent, and At ax an occasional, parasite. In a number 
of examples the nacre near the cardinal teeth was black and diseased. 
Z. rectus reaches quite a large size in this basin ; the largest example 
was 172 millimeters long and the dry shell weighed 375 grams. 

On account of its relative scarcity the value of this species as now 
occurring in the river would not amount to much. It is, however, the 
most valuable indigenous species so far as price per ton of shell is 
concerned. In luster it is not greatly inferior to L. anodontoides, the 
most valuable of the fresh-water mussels. Its elongate form makes 
it as well suited to the manufacture of knife handles as L. anodon- 
toides and for pocket knives the purple or pink nacred shells would 
not be so objectionable as in the manufacture of buttons, so that the 
advantage the other species has lies chiefly in the attractive color of 
its skin. At Grand Rapids we were told that a price of $40 per ton 
was offered for these shells. As this species lives under widely dif- 
ferent conditions, such as rivers, canals, and even lakes and ponds 
(though it is usually dwarfed in lakes), it would be one of the most, 
if not the most, promising species with which to attempt artificial 
propagation in the Maumee. 

29. Lampsilis ligament inns (Lamarck). Mucket. This is by all 
means the most common shell in the ]\Iaumee Basin, occurring so 
frequently in the river that it would be both needless and tedious to 
go into details. At Fort Wayne it was common in all three rivers — 
the St. Marys, St. Joseph, and the Maumee — and in the bed of the 
feeder canal. It formed the great mass of the 25 tons of shells col- 
lected by the clammers at the head of the Maumee. At Fort Wayne 
it reached its maximum development, and though not actually im- 
mense, it was quite large and heavy, as heavy indeed as could be 
worked up profitably. The shells decrease in size as we go down the 
river, so that those in the middle portion are of only medium size. 
Those in the Auglaize River were fairly large. Although L. liga- 
mentimis was found at most of the points examined for the whole 
length of the river, it became scarcer and showed a tendency to dis- 
appear from the lower stretches. It was not common in the second 
stretch of canal, though a few were seen in the canal bed below the 
dam at the very end of the trip. Only a small portion of the great 
number seen were taken ; however, we saved 282 examples. A number 
of the shells were measured and weighed, of which the largest was 



MUSSEL FAUNA OF MAUMEE RIVER. 53 

145 millimeters long and Aveighed 4G7 grams. The nacre of all the 
shells was white. 

L. ligamentinus^ the mucket, while not the very best of button 
shells, is not greatly inferior to the best, and on account of its com- 
monness and favorable shape and size is the most common species 
in commercial use. In the upper Mississippi it occasionally has to 
be discarded on account of its pink nacre, but this feature is wholly 
absent from Maumee shells. Perhaps 90 j3er cent of the shells of the 
Maumee Basin that possess conmaercial value are L. ligameiitinus. 

This species is rather free from parasites. A few Atax frequently 
occur, but Cotylas'pis^ Aspidogaster, and the other parasites found so 
frequently in the thin-shell mussels, are usually rare or absent here. 
Encysted distomids are fairly common, though hardly so common 
as in L. ventrlcosus. They are usually found about the edge of the 
mantle, though they may also be distributed throughout the body. 
The common form, designated in this report as the marginal-cyst 
distomid, is of especial interest in this species, as it is occasionally 
the cause of the formation of pearls. 

This species was not usually found gravid until late in the season. 
At the beginning of the Maumee investigation, in June, only a few 
examples were gravid, but the greater number w^ere barren. As we 
proceeded down the river, it was noticed that in the Auglaize River, 
August 8, the gills had begun to fill with glochidia and had a faint 
blackish edge. On a visit to the riffles of the St. Marys on September 
27 a large number of the mussels were found to be gravid. 

LamjmJ'ts ligamentinus is one of the most easily propagated of 
fresh- water mussels, and where it produces a heavy shell and white 
nacre, as it does in the Maumee, is one of the most valuable species 
for propagation. 

Although well-marked specimens of L. Ugamentinus are easily 
distinguished from the following species, L. luteolus, one frequently 
finds shells which are difficult to identify, apparently standing some- 
where between the two. 

30. Lampsilis luteolus (Lamarck). Fat mucket. Lampsilu lute- 
olus was found in about the same situations as L. ligamentinus but 
not in such abundance. Most of the examples seen were collected 
and the number of shells in our Maumee collection is 134. None of 
our specimens are of the variety rosaceous^ which is a form belonging 
to the St. Lawrence drainage. 

Unlike Z. ligamentinus^ this species is usually abundant in lakes, 
as well as being an inhabitant of rivers. It is indeed one of our 
characteristic lake shells. We obtained a number from Put-in Bay 
Island, Lake Erie, but they were rather small, as is usual with lake- 
dwelling individuals. The examples found in the St. Joseph River 
and the feeder canal were remarkably large and fine. This species 



54 MUSSEL FAUNA OF MAUMEE EIVER. 

appeared, like many of the others, to become smaller as we proceeded 
downstream. 

L. luteoUis is subject to infection by the same parasites as those 
which attack L. ligamentinus, viz, Atax and occasionally Cotylaspis. 
One example obtained in the mouth of the St. Joseph Kiver con- 
tained a number of Bucephalus polyTrwrphus. 

This shell has nacre of about the same quality as that of L. lig- 
amentinus^ but the shell is usually smaller, and has more lateral 
curvature, so that it can not be worked up so advantageously in 
making buttons. It is well adapted to the manufacture of fancy 
buckles, however. 

31. Lampsilis TnuUiradictus (Lea). This very attractive little 
mussel was not common in the Maumee Basin, and nearly all those 
found were dead shells, so nothing was learned concerning its habits. 
Only 9 examples were found, and these were in the upper two-thirds 
of the river basin, in the St. Marys, St. Joseph, and Maumee Rivers, 
and in the feeder canal. In 1907 a fair number were found in the 
feeder canal, and it had apparently thriven there to some extent. 
It is of no commercial importance. The species is a resident of lakes 
as well as rivers. 

32. Lampsllis ventncosus (Barnes). Pocketbook. This shell, 
though not especially common, occurs rather frequently in the Mau- 
mee Basin. At the riffles of the St. Marys 7 large examples were 
found. Very fine examples were also obtained in the feeder canal. 
In the reservoir of the feeder canal a fine but small specimen was 
found, pretty closely resembling L. capax. In the mouth of the St. 
Joseph River 18 were secured. Occasional examples were obtained 
all the way down the Maumee; in the Auglaize 14 fine specimens 
were taken. The largest ones found were in the Maumee below the 
dam at Defiance, and it was also common below the dam at Grand 
Rapids, and 2 more were secured in the Maumee above Water ville, 
Ohio. Altogether 83 examples were obtained. 

This shell also lives in lakes and 12 were obtained in Lake Erie 
at Put-in Bay, all dwarfed. 

Although well-marked specimens of this species are easily recog- 
nized, it has many deviations from the typical fonn. Some closely 
approach L. capax^ and we have seen shells from Lake Maxinkuckee 
which seem to lie between this species and L. Ivteolus. In other 
localities it appears to run into a species which in its typical form is 
known as L. ovatvs. However, in the Maumee Basin it is well 
marked and fairly uniform. 

Z. ventricosus is used to some extent in the button business, but it 
is quite frequently too thin for this purpose. 

In spawning season this is a very striking shell, the breeding 
mussel projecting long vividly colored flaps from the mantle and mov- 



MUSSEL FAUNA OF MAUMEE RIVER. 55 

ing" these with a spasmodic motion, while the animal lies on its back. 
The spawning season does not appear to be strictly confined to any 
particular season of the year, but spawning individuals can occasion- 
ally be found throughout the summer. One was observed in the St. 
Joseph Eiver at Fort Wayne on July 20. 

The principal parasites of L. ventricosus are Atax^ usually few in 
number, the distomid of Kelly which is frequently present but never 
numerous, and the marginal-cyst distomid, which is occasionally 
present in great numbers in the margins of the mantle and through- 
out the body. 

33. Micromya fahalis (Lea). This is the smallest and one of the 
most attractive of fresh-water mussels. It is rather rare in collec- 
tions, on account of its small size being easily overlooked. It was 
exceedingly abundant in the Feeder Canal; in 1908 we obtained a 
half dozen examples, and in 1909, by going over the ground thor- 
oughly in search of this particular species, several hundred were 
secured. 

34. Truncilla sulcata (Lea). Scattered valves of this species were 
seen along the shore of the St. Marys and Maumee, and now and 
then whole shells. Shells and valves were found on the shore of 
the St. Marys above Bluffton Street Bridge, along the St. Joseph 
near its mouth, and in the Auglaize River. Only 1 living example 
was found, and this was in the Auglaize. In all only 15 shells were 
obtained. 

Truncilla triqvstra was not found in the Maumee Basin, but we 
picked up several vales on the shore of Put-in Bay Island. 

GENERAL DISTRIBUTION OF SHELLS IN MAUMEE RIVER. 

In general, as to distribution of mussels in the Maumee River, cer- 
tain species were found to diminish in number or size, or both, in a 
downistream course. 

Qiiadrula hibercidata diminishes in both number and size until in 
the lower part of the river only a few dead and rather small shells 
were obtained. Dwarfed specimens were also found at Put-in Bay. 

Q. coccinea gradually diminishes and finally disappears at Grand 
Rapids, Ohio ; it was found at Put-in Bay. 

Q. ruhiginosa greatly diminishes in numbers, but a few remain 
until the very last. 

Q. cylindrical sparingly present around Fort AVayne, w^as not found 
far below Antwerp, Ohio ; no living specimens were obtained in the 
Maumee. 

PleurobeTMi clava diminishes in numbers and disappears at De- 
fiance, Ohio. 

Symphynota complanata was common and large in the feeder 
canal and is occasional in the St. Joseph at Fort Wayne. One speci- 



56 MUSSEL FAUNA OF MAUMEE ElVER. 

men was seen at New Haven, Incl. We ourselves found none in 
the Maumee, except at its very head. 

Ptychohrmichus phaseolus decreases in numbers and disappears at 
the very last near Miami, Ohio. Dwarf examples were found at 
Put-in Bay. 

Ohovaria c-ircidtos diminishes rapidly in numbers, and the last one 
was found in the Auglaize River at Defiance, Ohio. 

Lampsilis 7n idtiradiatus^ fairly well distributed in the upper part 
of the river, wholly disappeared at Defiance. 

L. ligaTnentinus decreased in number more than in size, but 
diminished considerably even in size. 

Trimcilla sulcata^ never very common, was not found below De- 
fiance, Ohio. 

The following species increase in number or size, or both, with the 
descent of the river: 

Quadinda pustidosa increases both in number and size until it is 
fairly common below Grand Rapids, Ohio. 

Q. lachrymosa first appears in the Auglaize a little above Defiance 
and becomes quite common in the lower part of the Maumee and in 
the canal. 

Unio gihhosus increases both in size and number and becomes one 
of the most common forms in the lower part of the river. 

Anodonta imhecdlis appeared in considerable numbers at the very 
last, in the canal below the dam at IMiami. Ohio. 

Ohliquaria refexa first appeared in the Auglaize River, near De- 
fiance, and became fairly common below there; the same may be said 
of Plaglola donaeiformis and PJagiola eJegans. 

Lmnpsilis gracilis shows considerable increase both in numbers and 
size. The same may be said of Lampsilis alatus^ of which very few 
were found at Fort Wayne, while it was abundant in the lower 
stretches of the river. 

The following species are of about equal size and distribution 
throughout the entire length of the river: 

Quadrula undulata was fairly common everywhere. It prefers a 
solid bottom overlaid with a thin stratum of rather soft mud, and 
wherever these conditions were encountered it Avas pretty sure to be 
found. 

Symphynota eostata was also fairly common everywhere; it in- 
creased somewhat in size as we descended the river, the examples 
found in the Auglaize being the largest obtained. 

A7iodonta grandis was nowhere abundant, but was found at nearly 
every station. It prefers a soft muddy bottom. 

Lampsilis iris was rare everywhere, but was found all along the 
river. 



MUSSEL FAUNA OF MAUMEE EIVER, 57 

L. rectus was nowhere found in abundance; the examples collected 
near Defiance were especially large and of fine quality. 

L. ventncosus was nowhere abundant, but was found occasionally 
all along the river. 

Of exceptionally rare shells, Plagiola securis was reprasented by 
only 1 specimen, obtained at the clammer's camp near Fort Wayne. 
In searching over the pile of shells found in the poultry yard referred 
to in the early part of this paper, it is believed that an example of 
Unio crassidens was seen, but as the shell does not appear in our 
collections its presence in the river can not be predicated witli 
certainty. 

In his paper on " The hydrographic basins of Indiana and their 
molluscan fauna," *» Dr. Call gives a list of 27 species of Unionicla; 
from the Maumee River; of these, 5 species, viz, Qnadrvla hci'o.s, 
Ahtsmidonta ealceola^ Ohovaria retiisa^ Ohovaria ellipsis, and Lanvp- 
silis subrostratus, were not seen by us. 

FOOD OF THE MAUMEE MUSSELS. 

The character of the food of mussels may be better appreciated 
after a short account of the method of feeding. 

If one looks at a mussel in its natural position in the water one 
of the first things to attract the attention is two openings, one 
large and usually conspicuously fringed with elongate papilla?, the 
other smaller and fringed or not, according to the species. These 
openings project from the posterior or sharp end of the mussel which 
is directed upward from the bottom. The larger fringed opening is 
toward the ventral portion of the mussel and is an incomplete tube 
formed by portions of the mantle placed together. This is the in- 
halent opening, and here water is taken into the gill chambers by 
means of waving cilia Avhich beset the gills, mantle, and other parts 
of the body bordering on the mantle chamber. The water thus 
brought in contains oxygen and food particles. The small cilia move 
the particles up to the anterior end of the body between the large 
labial palps or lips, which form a sort of funnel to the mouth. Here 
the food passes through the short gullet into the stomach, and thence 
into a long convoluted alimentary canal, which finally ends at the 
smaller of the two openings noted above, the exhalent aperture. 

There are minute pores in the gills, and through these the water 
is forced by cilia into the exhalent aperture, from which it returns 
to the lake or stream in which the mussel lives. Large robust mus- 
sels are able to produce quite a strong current, but this is usually not 
visible in the rippling water where they naturally live. It can be 
best observed in large river mussels removed from their native stream 



« Proceedings Indiana Academy of Sciences, 1896, pp- 248-257. 



58 MUSSEL FAUNA OP MAUMEE EIVER. 

and placed in the calm shallow waters of a lake. It is probable that 
these mussels breathe extra heavily in the lass highly oxygenated 
lake water. The current from the exhalent siphons boils up vio- 
lently and is ejected intermittently at more or less regular intervals. 

In many mussels there is no direct communication between the 
gill chamber into which the water, bearing solid particles, enters, 
and the suprabranchial chamber from which it is ejected through 
the exhalent aperture. Some mussels have the gills free above pos- 
teriorly, so that the two chambers have direct communication, and 
in these it is possible for the mussels to shunt solid particles directly 
from the gill chamber to the chamber above without passing through 
the alimentary canal. We have no evidence, however, that this is 
ever done. The mussel probably exercises little choice as to the 
nature of its food. The papilla? which fringe the edge of the in- 
lialent orifice are supposed to be water-testing organs, and the mussel 
can close itself against foul water or against positively objectionable 
material. In the feeding mussel, as observed in the stream or lake, 
there is no appearance of nicety or fastidiousness as to the particles 
swept in ; the wide open, immobile, and expressionless lips admit the 
general current and its diverse minute population with a catholic 
impartiality. The alimentary canal of the mussel is usually filled 
with a sort of impalpable or sometimes gritty mud, and mixed in 
Avith this are found scattered organisms of various sorts which serve 
as food. 

The impalpable mud usually corresponds closely in color and 
general appearance with the bottom in which the mussels are found ; 
in rivers with yellow clay bottoms it is yellowish; in dark, muddy 
bottoms it is dark. It consists of fine particles which have been 
held in suspension in the water. In turbid streams it consists of 
the material which gives the water its turbidity, in clear ponds 
with bottoms of fine mucky silt it is black, and in those still waters 
full of diffused minute alga? and flagellates, which are common in 
ponds and pools, it is predominantly organic and of a greenish cast. 
In the latter case it usually consists almost entirely of colonies of 
plankton-scum (water bloom) alga^, which are frequently inclosed in 
clear vacnolelike spaces. 

The moderately long alimentary canal suggests that the ingested 
material contains a low amount of nutriment; observations also seem 
to indicate that the digestive powers of the mussel are slow and feeble. 
In mussels which had been kept for several days ScenedesTrms was 
found as fresh and green as ever, and in one mussel obtained in the 
reservoir of the feeder canal green flagellates were still alive and ac- 
tive after having been at least 36 hours in the mussel. While in these 
instances it is possible that digestion was delayed by the capture of 
the mussel and placing in changed surroundings, there are other indi- 



MUSSEL, FAUNA OF MAUMEE EIVER. 59 

cations that digestion is normally slow and imperfect. In mussels 
taken from the beds, washed, and immediately examined, diatoms 
found at the posterior end of the alimentary canal are frequently 
alive and exceedingly active ; they appear to be even more active than 
when taken from open water, and the form and color of the chromato- 
phores are unchanged. It is possible that in these cases the mussel 
obtains some nutriment from the outer coating of the diatom and that 
the removal of this makes the latter especially active. 

The organisms found in the alimentary canal of mussels vary con- 
siderably with the places in which the mussels live, but, so far as 
observed, do not differ much, if at all, among the various species. 
They consist of such planktonts as are small enough to be taken in, 
and not active enough to resist the force of the inward current pro- 
duced by the mussel. The largest and most active planktont taken 
to any extent was the entomostracan Bosminia. This was found 
with considerable frequency in lake-dwelling mussels, but as all seen 
were dead, and many were simply empty shells, it was not possible 
to ascertain wdiether the mussel had taken in the living animals or 
not. Actively moving flagellates are quite frequently ingested. In 
the gill chamber of some Quadrula tiihercnlata^" a species with an 
exceptionally large inhalent opening, we found such objects as Chiro- 
nornv,^ larvae and one small mature dipterous insect. 

There are several organisms which are likely to be found in mus- 
sels wherever taken. Species of Scenedesmus are almost invariably 
present, as is also Navicula along with other diatoms and species of 
Pediastrum^ such as P. pertuswn with great frequency, P. horymvum 
rather commonly, and P. ehrenhergii^ which appears to be a rare 
species everywhere, only occasionally. The rotifer Anurma cochlearis 
is also found frequently and under all sorts of conditions, in lakes, 
ponds, and rivers. The part usually found is the empty lorica. 
Another common object found in the alimentary tract of mussels is a 
club-shaped, several-celled, brown object, probably the spore of a 
fungus or lichen. 

With the exception of those just mentioned the organisms found 
within mussels are very diverse, and the creatures seem to be quite 
omnivorous within the limits of what they can capture. Just as in 
any stream one finds various types of planldon aggregates from the 
extreme type of " potamoplankton " or characteristic river plankton 
of the mid-current, recognizable by the scarcity of water-bloom alga?, 
to well-developed types of " limnoplankton " or lake plankton in the 
quiet sloughs and bayous, so in these different portions of the stream 
one finds different sets of organisms in the mussels. In lakes full of 
diffused water-bloom algse, the material which forms the mudlike 
matrix of the food in river mussels is to some extent replaced by the 

« Not in the Mauniee River, however, where few were examined. 



60 MUSSEL FAUNA OF MAUMEE RIVER. 

water-bloom algse. These may serve also as food. The stomach con- 
tents of such mussels, when placed in vials of preservative, differen- 
tiate into a lower stratum of black mud and an upper flocculent 
stratum of fairly pure algse. 

The mussel may be regarded as a sort of living filter, feeding upon 
the filtrate it separates from the water. It would be worth investi- 
gating to ascertain whether they arrest and destroy pathogenic 
bacteria and thus become efficient purifiers of water. They are valu- 
able aids in plankton investigations, capturing many minute organ- 
isms which escape methods of collection ordinarily in use. Though 
at the outset it hardly looks as if they could cope with the problem, 
their capacity for ingesting water-bloom algse {Clathrocystis, 
Lynghya^ etc.) suggests that it might be worth while to investigate 
their efficiency as reducers of plankton scum. 

The Avater bloom, as is well known, frequently becomes a nuisance. 
It collects in. noisome masses of scum along shores of otherwise at- 
tractive lakes and reservoirs. Its taste and odor renders the water 
of reservoirs undrinkable, and for a remedy of this condition the 
copper sulphate treatment has been devised. Its presence in large 
quantities along the shores of our most beautiful lakes renders them 
unattractive during the late summer season, and in many places 
brings the bathing season to an early close. Aside from the filthy 
appearance of the w^ater, many persons claim to be actually poisoned 
by the water bloom, and there are instances on record of live stock 
being fatally poisoned by drinking water covered with plankton 
scum. In addition to this, the great amount of decaying mate- 
rial in the water is said to take up oxygen, making the lower 
strata iminhabitable for fish. It is possible that by planting large 
numbers of mussels, supplemented by planting ^^g masses of Ghiro- 
nomns, which appear to hatch easily and the larvae of which eat the 
coarser alga?, that the plankton-scum nuisance can be gTeatly abated. 

The stomach contents of mussels taken from the main current of 
the St. Marys, St. Joseph, and Maumee Rivers were rather note- 
worthy for their paucity of organic material. Through the large 
mass of muddy matrix filling the stomach were usually scattered a 
few Scenedesmus, various diatoms, and an occasional Pediastrum or 
Cosmanum. At the riffles small brown cystlike objects, which may 
have been a species of Trachelcmonas, were quite common ; with the 
exception of this the mussel contained very little. Among the organ- 
isms noted were Seenedesmus cmidatus^ Cmlastrum microsforum^ 
Pleurosigma, several forms of Nmncula^ PJiacus longicaudus^ Pe- 
diastrum haryanum^ Gomjjhonema^ a sponge spicule, and an active 
Euglena-Wko, organism. The stomach contents of mussels talvcn else- 
where along the river and in its tributaries, as in the Auglaize at 
Defiance and the Maumee at Grand Rapids, were not essentially dif- 



MUSSEL FAUNA OF MAUMEE KIVER. 61 

ferent. One of the examples of L. ligarrhentinus taken in the 
Auglaize, however, contained many very minute cylindrical objects 
with rounded ends, which were probably bacteria. 

The stomach contents of the mussels found in the reservoir of the 
feeder canal, a shallow pondlike body of water, bore considerable con- 
trast to that of the river. This reservoir was well populated with 
small gi-een monads and similar organisms, and during the last days 
of our observation became covered over Avith a green scum. It was 
" puddle plankton " rather than that which is characteristic of either 
lakes or rivers. The stomach contents of the nnissels were gTeenish 
in mass, and were found to consist of various organisms, mostly green 
in color and actively moving. 

The main mass consisted of small globular and thick celled, green 
or brownish flagellates, probably T raehelernonas lagenella. Among 
other organisms were species of Phacus^ several specias of Scene- 
desmus^ Pediastrwm lyertuHum^ GonvphoTiema^ several species of 
Navicula^ a little Botryococcus brwini^ Anunea cochZearls^ Cos- 
marium, small fragments of a Conferva-Vike alga two or three cells 
long, fragments of the test of Ceratium hirundinella, and the brown 
objects resembling fungus spores. There were numerous narrow 
curved objects which were probably loricas of T raehelernonas. 

PARASITES OF THE MAUMEE MUSSELS. 

During the progress of the Maumee lliver investigations the para- 
sites encountered were noted and examples saved for further study. 

Generally speaking, Anodontas and other thin-shelled mussels are 
more heavily parasitized than the species having thick shells, and 
parasites are more abundant in shallow, warm, and quiet pools than 
in rivers. 

The mussels of the feeder canal reservoir were more heavily para- 
sitized than those of the main streams. The following parasites were 
observed : 

1. Cotylasjns indgms Leidy, a small organism, which to the naked 
eye resembles a pale leech. The body is trumpet shaped, and the 
ventral surface has a large ovate disk cut up by partitions. By this 
disk the animal adheres to its host, and it usually inhabits the axils 
of the inner gills. It was most common in Anodonta gmndis and 
occurred occasionally in most other species of mussels examined. An 
examjole of Lampsilw rectus obtained at the riffles contained the sur- 
prisingly large number of 67 of these parasites. So far as could be 
ascertained, this parasite produces no marked physiological effect 
upon the host and appears to do no particular harm. 

2. Aspidogaster concMcola Von Baer bears a close general resem- 
blance to the above, but is considerably larger, and the adhering 



62 MUSSEL FAUNA OF MAUMEE EIVER. 

disk, which is hirger and more complicated, has been aptly compared 
to the sucking disk of Remora. 

This parasite affects various species of mussels, but its favorite 
hosts are species of Anodonta, Synvphynota complanata,, and Lamp- 
s'dis alahfs, in which considerable numbers were found. It usually 
inhabits the pericardial cavity of the host, though when considerable 
numbers are present they may be found in other parts of the body. 

The finding of a small elongate " hinge pearl " in a mussel affected 
by these parasites suggested that, under peculiar circumstances, the 
bodj^ of one of these creatures may form the nucleus of such a pearl, 
since the shape of the two is quite similar. This supposition, however, 
needs further investigation. Aspidogaster reproduces within the 
jDericardial cavity of the host without any marked metamorphosis, 
and they are found of all sizes, the minute ones having the same form 
as the adults." 

3. The onarginfd-cyst disfomid. — A distomid forming spherical 
cysts was fairly common, especially in Lampsilis Ugamentimis and 
L. ventricostis, along the whole length of the river. A few of these 
parasites were also found in one Quadrula vndvlata and one Sym- 
pliynota costata. It is probably the species discovered and briefly 
described by H. M. Kelly,* who noticed it in four examples of L. 
ligamentimis. 

The cA'sts are usually found along the edge of the mantle, generally 
in the muscular portion below the pallial line. They are also fre- 
quently embedded in other parts of the body, such as the adductor 
muscles, and esjDecially in the keel of the foot, where they are occa- 
sionally found in great numbers. There are indications that they 
prefer muscular tissue. 

Cysts of various ages were frequently present side by side in the 
same mussels. The youngest cysts are translucent and refractive, 
faintly yellowish in color, and resemble minute pearls embedded in 
the edge of the mantle. As they grow older they form conspicuous 
black dots. One of the smaller cysts measured was 0.3 millimeter 
in diameter, and a large blackish opaque one was 0.9 millimeter. 

Examined with a microscope the younger cysts appear as brick-red 
spheres, crossed and recrossed by irregular cracks like those of a dried 
mud flat. Upon carefully breaking open this crust a minute distomid 
is released, which slowly crawls about on the slide. One of the dis- 
tomids thus released was colorless, 0.365 millimeter long and 0.21 
millimeter wide at its widest portion. Viewed from above, it was 
pear shaped in outline, the anterior end being acute, the body gradu- 
ally broadening behind and the posterior end being rounded. It had 

" The development of this species is given by Huxley, Anatomy of Invertebrate Animals, 
p. 178. 

* Bulletin Illinois State Laboratory of Natural History, vol. v, p. 40G. 



MUSSEL FAUNA OF MAUMEE EIVER. 63 

tv.o suckers, an anterior one at the small anterior end and a ventral 
one, about the same size, placed far back near the swollen posterior 
end. The posterior part of the distomid contained a mass of black 
material, probably food. 

As the C3^st increases in age it becomes black and forms a conspic- 
uous dot about the size of a pinhead in the tissue. It is found on 
examination that the cyst itself, which is surrounded by the black 
material, has not increased in size, and the wall retains its brick-red 
color. The appearances indicate that the black material, which is 
probably a product of excretion of the parasite, has been passed 
through the cyst wall. 

In our alcoholic material the cysts, which appear as small black 
spots to the naked eye, show under the simple lens as minute orange 
globes, surrounded by a semitranslucent halo, the alcohol having 
apparently dissolved out the black material around the cyst wall. 

The marginal-cyst distomid reminds one in its general appearance 
of Diplostomum^ which is frequentl}'' found encysted in the skin of 
fishes, especiall}^ minnows, forming small black spots. In the fishes, 
however, the black color surrounding the cyst appears to be due to a 
deposit of pigment by the fish. 

The effect of this parasite (the marginal-cyst distomid) upon the 
mussel appears to be quite various, according to the location of 
the cyst. In many cases, probably where the cysts are near the 
inner surface of the mantle, no effect whatever appears to be pro- 
duced upon the shell. Under other cysts the shell is stained brown, 
and where the cysts are near the outer.surface of the mantle the whole 
edge of the shell is blistered irregularly and stained a steel-blue color. 
Moreover, as the cysts increase in size they form firm nodules in the 
mantle and cause the shell to be built out around them, so that they 
leave pits in the nacre similar to those left by pearls. 

In studying the collection of shells of L. Ugamentinvs at the 
Bureau of Fisheries, which contains perhaps 2,000 specimens of this 
species and is represented by shells from widely separate parts of the 
country, peculiar malformations of the shell were observed, which 
were probably caused by the parasite under discussion. Among these 
markings are pits near the margin of the shell, such as we have 
frequently observed in the living mussel underneath the cyst, and 
sometimes these contain small attached pearls. One of the most 
marked features is a tendency of the antero- ventral portion of the 
shell to cease growing outwardly and either become thickened or 
gape away from the middle line so that the mussel can not tightly 
close its shell in front. It appears that the presence of the cysts 
keeps the affected portion of the mantle in a constant state of con- 
traction, thus preventing it from building the shell outward in the 

22367°— 12 5 



64 MUSSEL FAUNA OF MAUMEE EIVEE. 

normal fashion. In some of our alcoholic material this contraction 
and thickening- of the mantle is quite markedly shown. The effect 
of this peculiar form of growth on the shell as a whole is to give it a 
somewhat more inflated and elongate appearance than is character- 
istic of the species. Shells of L. Ugamentinus thus affected approach 
the contour of L. luteohi8\ and while close approximation of these 
two species is not rare in normal shells, some of the puzzling cases 
are probably L. ligameyitiniis which has been modified by this dis- 
tomid. 

In L. ventiicosKS^ which is ver}^ susceptible to the attacks of this 
parasite, we frequently have gaping shells, and others which are 
markedl}^ thickened; Some of these thickened shells closely resemble 
those of L. ligmnentinus. The marginal-cyst distomid is of especial 
interest on account of its connection with pearl formation. 

Suspicion that this distomid had some connection with the forma- 
tion of pearls was aroused when the objects were first encountered, 
as their form and position was especiallj^ suggestive. This suspicion 
was intestified by frequently finding small round pearls in the mantle 
of mussels, closely associated with the cysts. Finally, on decalcifying 
and clearing in paraffin and x^dol, preparatory to embedding, a por- 
tion of the mantle containing a small round free pearl surrounded by 
the cysts, a c^yst could be clearly distinguished in the center of the 
pearl. Our material, which was preserved by simply dropping the 
mussels into strong alcohol, was not in good condition for histological 
jDurposes, so that we were unable to demonstrate the nature of the 
tissues surrounding the pearls. 

Our studies indicate that the marginal-cyst distomid is widely dis- 
tributed. "We have found the cysts in alcoholic material collected in 
the Mississippi River, in L. Ugamentinus collected by Freeland and 
Williams in the Illinois River in 1907, in L. ventricosus collected in 
the Yellow River, Ind., by ourselves in 1008, and in L. ligam£ntinus 
collected in the Ohio by Boepple in 1009. From the appearance of 
shells examined, we judge it to be common in the Minnesota River. 

4. The distomid of Oshorn. — Quite frequently during our mussel 
investigations we have found Anodonta affected with small white 
sporocysts, which covered the outside surface of the mantle next the 
nacre. The nacre of these mussels was often raised into a number of 
sharp pearly blisters, and in some cases small white areas, conform- 
ing with the size and shape of the sporocysts, indicate that the sporo- 
cysts had been actually covered with nacre. Mussels thus affected 
were found in the Little River ditch near Aboite Station, Ind., and 
in White Bear Lake and Lake Minnetonka, near St. Paul, Minn., 
in 1008. Notes on alcoholic examples of Anodonta collected in AVild 
Cat Creek, Carroll County, Ind., in 1809 refer to '' white distomids," 
which were probably this species. 



MUSSEL FAUNA OF MAUMEE lUVER. 65 

111 the reservoir of the feeder canal at Fort Wayne we found an 
Anodonta plentifully infected with these sporocysts, and were able 
to make out in detail some of the steps in the life history of the 
animal. 

The sporocysts were small white objects, elliptical in outline, about 
1.14 millimeters in leng-th and 0.05 millimeter w4de. The skin of the 
sporocysts was transparent enough to reveal, in the interior, the cer- 
caria>, which were three in number in most cases examined, closely 
doubled up and snugly packed together, and performing an uneasy 
wiggling motion. The walls of the sporocyst easily rupture; in an 
example studied at St. Paul the previous year the covering was torn 
apart by a dissecting needle and the cercarian set free. In the ex- 
ample now under discussion the sporocysts were rupturing of their 
own accord, allowing the cercaria? to escape. The following is a de- 
scription of the various stages observed : 

Cercaria just escaped from the sporocj'st: 

Body elongate cylindrical with bluntly rounded ends and divided 
by a well-defined constriction into an anterior two-thirds, the body 
proper, and a posterior one-third, the ^ tail.'' Body portion ellip- 
tical in outline with the two ends similar. Anterior sucker terminal, 
about one-fourth the greatest body diameter, circular, with concen- 
tric ridges faintly marked. 

Posterior sucker one-third body diameter, also circular, but with 
radiating ridges, situated almost in the exact center longitudinally 
as well as transversely; constriction only one-seventh the bodv diam- 
eter with a well-defined groove all the way around. Posterior por- 
tion (tail) almost perfectly elliptical, twice as long as wide, both ends 
alike and more bluntly rounded than the body proper. Tail covered 
with a thick epidermis, raised into longitudinal ridges close together, 
very narrow and zigzag in arrangement. 

The internal structure is indistinct, but far enough developed to 
distinguish the pharynx and digestive canals. The former is rela- 
tively smaller than in the adult and is much nearer the posterior 
sucker than the anterior one. This means that in after development 
the posterior sucker travels backward, while the pharynx travels 
forward. At this stage the whole interior of the body is filled with 
large cells having numerous and interlacing intercellular spaces. 

After remaining under the cover glass for a little time the tails 
break off at the constriction and the body moves about alone. 

The following is a description after the tail is shed : 

Body elongate, ovate, without any traces of segmentation or sepa- 
ration into parts; anterior end evenly rounded, posterior end drawn 
out into a point, bluntly rounded at the tip. 

Anterior sucker terminal, circular, three-sevenths the diameter 
of the body, with circular ridges. Posterior sucker on the mid line 



66 MUSSEL FAUNA OF MAUMEE EIVER. 

of the ventral surface, four-sevenths of the entire length from the 
anterior end, circular, with radiating ridges and four-sevenths the 
diameter of the body. Posterior end of body with traces of the zig- 
zag longitudinal ridges seen on the tail of the young. Mouth in the 
center of the anterior sucker, opening into a spherical or transversely 
elliptical pharynx close behind the sucker and one-third its diameter. 
From the pharynx two digestive tubes lead back to the posterior end 
of the body, one on each side, each a little narrower than the phar- 
ynx, of nearly uniform diameter throughout, and curved outward 
parallel to the lateral body margin. They reach nearly to the poste- 
rior end of the body, where each ends in a network of intercellular 
spaces and tubes, the excretory organ. 

The newly escaped cercarium is about 1.05 millimeters in total 
length and 0.2 millimeter wide. The length and width are con- 
stantly changing, according as the organism is stretched out or 
contracted. 

The tail is no longer an organ of locomotion; it is very large and 
unwieldy and easily detached ; most of the cercaria shed it soon after 
emergence from the sporocyst, so tliat it is evidently a vestigial organ. 

After shedding the tail the organism, which now has the typical 
form of a Distomicm, moves up to the umbonal region of the mussel 
and acquires a pinkish color. From the umbonea it spreads in di- 
verging lines postero-ventrally and antero-ventrally and finally fills 
in the space between. "VVliere it is located it profoundly affects the 
nacre, staining it a salmon color, and frequently giving it a diseased 
appearance. The presence of this distomid in Anodonta and its re- 
lation to the salmon-colored nacre was first pointed out by H. L. 
Osborn" and further discussed by H. M. Kelly.'' 

In its general appearance and in its life history this organism 
agrees very closely with Dlstomum dupllcatwn as described by von 
Baer, who gives a detailed description of the sporocyst and a history 
of the escajDe of the cercaria and its subsequent behavior.*' It differis 
from that species in the fact that the tail is not clavate, is longer 
than the body, straight, and not pediculate. It differs from the 
cercarian genus Rhophalocerca Diesing, formed to include D. dupli- 
eatu?n; only in the fact that the tail is not clavate. Von Baer de- 
scribes what appeared to him a form of reproduction in this distomid, 
but from what is now known of related forms it appears probable 
that he misinterpreted appearances. Dhtomum duplicatum. (known 
also as Rhophalocerca tardigrada Diesing) is of special interest as 
the organism first shown by De Filippi to be the cause of pearls in 



« Zoological Bulletin, vol. \, no. 6, 1898. 

» Bulletin Illinois State Laboratory Natural History, vol. v^ art. viii. 
<■ Nova Acta Physico-Medica Academia? Ctesares Leopoldino-Caroliuce, Natural Curio- 
eorum, vol. xiii, pt. 2, 1827. 



MUSSEL FAUNA OF MAUMEE RIVEK. 67 

Anodonta cygnea of Europe; hence a few remarks concerning it may 
not be out of place. 

So far D. (luplicatum lias not been reported from any mussels ex- 
cept Anodonta. Its discoverer found it usually in the kidneys of 
Anodonta ventrtvo.lia and more rarely in the same organs in A. ana- 
tliui. One badly infected example of this latter species had the 
sporocysts in the kidneys, gills, pericardial cavity, and edge of the 
mantle, and in an example of .1. veiitncosa they were even present in 
the general circulation. Filippi found the sporocysts and free dis- 
tomids in the mantle of A. cygnea^ some of them surrounded by nacre ; 
and Pagenstccher found them in the tissue of the genital glands, 
liver, and kidneys of the same species. In this country Leidy° has 
reported ^'■Rhophalocerca tardigroda attached to the mantle of Ano- 
donta fwoiatilis; specimens also obtained by Mr. Lea from A. 
lacKstr'isy 

The mature form of Dhtomnni dv/plicatum is said to be D. tereti- 
colle Rudolphi, a parasite of fresh-water fishes. 

To return to the distomid of Osborn: In addition to its being 
found in Anodonta grandis, this parasite (in the distomid form) has 
also been reported from Strophitus edentidus by Osborn. We had 
very few living mussels of this species to examine; however, we 
found dead shells with the peculiar salmon-colored nacre, similar to 
that produced by this parasite in Anodonta. 

In vicAv of the fact that a Distomum quite similar in appearance 
lo this is found in other species of mussels, while sporocysts are rare 
(we found them only in the liver tissue of one example of Qumlnda 
tiiherculata in Tippecanoe River), it becomes an interesting ques- 
tion whether this distomid migi-ates from the Anodonta to other 
mussels after issuing from the sporocyst. In the first place, such a 
migration is probably not an event closely connected with the life 
history of the species; for the tail, which functions as an organ of 
locomotion in those species which do migrate, has here been allowed 
to become functionless, or even a dead weight. Moreover, if the host 
of the mature form is a fish, as one might expect it to be from the 
life history of its near relative D. dupUcatmn,., it would be to the 
advantage of the distomid to remain in the thin-shelled Anodonta, 
which is much more likely to fall a prey to fishes, or indeed any 
other predaceous animal, than the thicker-shelled species of mussels. 
It is very likely, however, that, just as in greatly infested mussels 
the cercarise frequently overflow the organs which appear to be their 
favorite habitat, so a considerable number of distomids may be 
crowded out of the host mussel and be forced to seek protection and 
food elsewhere. According to von Baer ^ the sporocj^sts of Distomum 

« Proceedings of the Academy Natural Sciences, Philadelpliia, 1858, p. 110. 
'' Loc. cit. 



68 MUSSEL FAUNA OF MAUMEE RIVER. 

dwplicatum' and their contained ceicariro live for a considerable time 
in the water when removed from the host. Migration wonld there- 
fore be possible. 

It has so happened that the Anodontas in which we found sporo- 
cj'sts were all taken in early and middle sunnner, in Jul}- or August; 
we found free distomids, hoAvever, as early as work in the field began, 
in June. It would require the study of infected regions throughout 
the year to ascertain whether the various phases in the life cycle of 
this distomid bear any relation to the seasons. Von Baer found his 
Anodontas affected by the sporocysts of D. dupUcatum in late autumn. 

5. The distomid of Kelly. — In several of the thicker-shelled species 
of mussels, such as Lampsilis ventHcosus, L. Uffamentinus, Ohovaria 
chxulus, and especially in Qu-adrula tuhercidata and U7uo ffibhosus, 
one frequently encounters pink distomids bearing a close similarity 
to the distomid of Osborn mentioned above. Kelley," who first de- 
scribed and discussed these distomids, was of the opinion that they 
are identical with those described by Osborn as affecting Anodonta 
and Strophitus and producing a discoloration of the nacre. 

Indications pointing to the identity of these two distomids are, 
first, the fact that dorsal baroques are formed in Anodonta by the 
distomid of Osborn, and second, that the distomid of Kelly, in addi- 
tion to forming baroques in Ohovaiia circtdus, frequently causes the 
nacre to become rough and assume a yellowish color, approximating 
the salmon color of affected Anodontas. 

In their typical form as they appear in the thick-shelled species of 
mussels, these distomids exhibit several minor points of difference 
from the distomid of Osborn. Practically they present so different 
a set of manifestations that in our field notes it was found necessary 
to separate them. 

After considerable study and comparison we are inclined to think 
them the same, but in this discussion, since the point of view is 
ecological rather than taxonomic, it will conduce to clearness to refer 
to this manifestation of the species under a different name. Simi- 
larly, it is convenient and customary to speak of the cercaria of a 
distomid, the nanplius of a crustacean or the leptocephalus of an eel. 
The distomid of Kelly is usually found in much fewer numbers than 
that of Osborn, a fact which suggests that it may be erratic or 
a stray migrant, and so far we have not found it associated with 
sporocysts or cercaripe except in the single instance of Quadrula 
tuhercKlata referred to above. It is usually confined to the outer 
surface of the mantle near the cardinal or lateral teeth of the mussels 
affected, and it is frequently associated, as Kelly has pointed out, 
with irregular dorsal baroques or pearls. It does not produce a 
salmon-colored stain in the nacre of the mussel affected, and usually 

« Bulletin Illinois State Laboratory Natural History, vol. v, art. 8. 



MUSSEL FAUNA OF MAUMEE EIVEE. 69 

produces no stain at all. The discolorations frequently found ni the 
umbones of mussel shells can not all be attribut(^d to it, since in one 
pond where such stains are very frequent, no distoniids could be 
found. In general, it is a baroque former rather than a stain pro- 
ducer. In some localities it especially attacks Unio g/hhosus and 
Qvadrula ttiherculata^ and the host mussels are peculiarly prolific 
in baroques. Most of the dorsal baroques which the clammers save 
and sell to dealers in jewelry are probably formed through the ac- 
tivities of this parasite. The baroques are known among the clam- 
mers as " chicken feed " and usually bring about $2 per ounce on the 
ground. In many places this material is fairly common and is a 
source of revenue among the clammers. 

It is probable that the distomids which stray into large heavy- 
shelled mollusks fall out of the normal course of their life history 
unless they again migrate to thinner-shelled species, for the thick- 
shelled mussels, after they have grown, are proof against fishes, musk- 
rats, or any other predaceous animals, and live to die a natural 
death, or in these recent times to fall a prey to the clammer. This 
gives the parasitic distomid no opportunity to enter into another 
host in the usual manner. Crawfishes eat mussels that have died ; 
but so far as known the host of the mature distomid is always a 
vertebrate. 

Moreover, on the supposition that the distomids found in the 
heavy-shelled mollusks are migrants from Aiiodonta, SfrophUxs, or 
other thin-shelled species, these mussels, which are regarded as 
valueless by the clammer and destroyed by the wholesale, are of im- 
portance as an intermediary in the formation of baroques and perhaps 
all distomid-formed pearls. Similarly, though in a somewhat dif- 
ferent manner, the cockle, Cardii/m, acting as an intermediate host 
between a species of mussel-eating duck and the salt-water mussel 
Mytilus, brings about pearl formation in the latter species, as shown 
in the investigations by Jameson « of pearl formation in Mytilus. 

6. Bucejjlialus polymorphiis (von Baer). — During the Maumee in- 
vestigations this species was encountered in only one instance, when 
an example of Lampsilis luteolus. obtained near the mouth of the St. 
Joseph River, was found to be a fleeted. In investigations outside 
the Maumee Basin it was not infrequently met. According to Kelly, 
who has had considerable experience with mussel parasites, it is 
fairly common, though not so common as Aspidogastei^^ Atax, or 
Conchopthirus, and affects a large number of species of mussels. 

Bucephalus pohjmorphus is of considerable economic significance, 
since it has been shown by von Baer and later b}^ Kelly tha*: this 
species (and, as Kelly has pointed out, some other distomids as well) 
frequently affects the generative tissue of the host mussel to such an 

"Proceedings Zoological Society London, vol. \, 1902. pt. 2, p. 140-166. 



70 MUSSEL FAUNA OF MAUMEE RIVEK. 

extent as to wholly obliterate it and render the mussel incapable of 
reproduction. In those mussels {Lampsilis gracilis, L. luteolus, etc.) 
in which there is a marked difl'erence in the form of the shell of the 
two sexes, Bucephalus, by arresting sexual development at various 
jDeriods of the mussel's life, stops also the development of the peculiar 
form of shell characteristic of the sex. Shells are thus left at vari- 
ous intermediate stages of sexual development, though they may 
attain full size, and this has doubtless led to considerable perplexity 
in identifying them, or even to the descriptions of new species. It 
may also be partly responsible for the large amount of synonymy 
found in the Lampsilis rentricosus group. 

It can be readily inferred that the introduction of BucepJmlu^- 
infected mussels into places where breeding or breeding experiments 
are being carried on would be likely to produce disastrous results. In 
collecting mussels for propagating purposes, therefore, shells of in- 
different contour as regards sexual peculiarities should be viewed 
with suspicion. In addition, examples should be dissected to detect 
signs of the presence or al^sence of the trematode. 

The mature form of Bucephalus polymorphus is Gasterostonium 
fnhhriatunv von Siebold, which is a parasite of the pike {Esox) and 
perch {Perca), and an intermediate host (eaten by the final host) is 
some species of minnow. {Leuciscus is one of the knoAvn hosts.) In 
ponds devoted to the breeding of mussels the ravages of Bucephalus 
might be held in check by avoiding the peculiar combination of mus- 
sel, minnow, and pike or perch, provided of course these were not the 
particular species of fishes required to carry the glochidia. 

7. Af<cans-\\\s.Q worms Avere occasionally fomid in the alimentary 
canals of mussels, and are probably to be regarded as parasites. 

8. Leeches were found quite frequently on the inside of the shells 
of Anodonta, in all probability genuine parasites. They appear to 
be quite fond of the mucus of mussels, as is shown by the following 
observation: On the inside of a number of valves cleaned by musk- 
rats and left on the bottom in Lake Maxinkuckee, the leeches were 
found to have accumulated in considerable numbers. Leeches also 
frequently attack other species of mollusks. as Planorhis, in large 
numbers. It could not be ascertained whether they injured the mus- 
sels to any marked extent or not. 

9. Atax ypsiJophorvs was a common parasite of mussels through- 
out the length of the river, and was particularly common on Ano- 
donta in the reservoir of the feeder canal. An example of Ataos, 
probably this species, found in a mussel at the riffles of the St. Marys 
River, was so heavily burdened with a growth of Vorticella on its 
body that it could hardly move about. We came across several other 
specimens similarly affected. 



MUSSEL FAUNA OF MAUMEE EIVER. 71 

Atax intermedius and Atdx tumidus were also common ou the 
mussels in the reservoir. There were probably several other species 
of Atax present; a small species with a red spot posteriorly was quite 
common. Lampsilis alatus below the dam at Grand Kapids was 
particularly heavily infected; in three examples 290 mites were 
found, thus averaging about 97 per individual. 

Many of the Atax were full of eggs, and eggs and larvas in differ- 
ent stages of development were common but not markedly abundant 
on the inner side of the mantle and in the gills of the mussels. We 
did not observe this 3^ear what was so conspicuous a feature in 1907 
of the mussels found in the lakes about St. Paul, Minn., at the same 
season of the year, viz, the young mites clustered thickly about the 
papillae of the inhalent siphons. The mites were apparently not 
breeding as heavily in the Maumee, and the young were not so far 
advanced in development at the date examined. Atax probably pro- 
duces considerable irritation in the mussel, but we have no evidence 
that it atfects shell characters. "Where they bury their eggs in the 
tissue of the mussel they leave small but conspicuous scars which are 
apparently permanent. 

Although it has been suggested that the eggs of Atax may lead to 
the formation of pearls, we have been unable so far to find con- 
vincing evidence on this point. In our studies of mussels of the 
Wabash Basin we obtained a great number of calcospha'ric struc- 
tures, many of them in situations that suggest a close relation to Atax 
eggs. This subject is reserved for future investigation, as it does 
not pertain to Maumee studies and would lead too far afield. 

AYherever we have examined mussels we have encountered the 
parasite Concho pthhnis^ a protozoan bearing considerable superficial 
resemblance to Paramoxium. This organism is of such universal 
occurrence that we have not taken trouble to specifically mention its 
presence. There are several species {curtus, anodonta^, etc.) para- 
sitic on fresh-water mussels. 

GENERAL CONCLUSIONS. 

The IMaumee River, as it exists to-day, is not on the whole a 
stream where clamming operations could be carried on profitably, 
as it would very quickly become exhausted if such operations were 
undertaken on a large scale. The mussels were apparentlj^ once much 
more numerous than at present. There are two conditions which 
militate against the well-being of mussels in the upper part of the 
river: The large amount of sewage, especially refuse from the gas 
works, and the irregularity of flow of the river, which has been in- 
creased by the removal of much of the original forest along the river 
•banks. The best mussel beds in the river are at Fort Wayne, Ind., 
and below the dam at Defiance, Ohio. By far the greater number 



72 musseIj fauna of maumee river. 

of marketable shells in the river are Lcwipsilis Jigcnnentinus, and the 
most valuable native species is L. 'rectus. 

The species of mussels to be recommended for propagation natur- 
ally varies with different rivers. In the Maumee, the species gener- 
ally regarded as the best button shell, the niggerhead, Quadrula 
ehena., could not be recommended as it is doubtful whether it would 
thrive in the shallow water. On the other hand, the mucket, Lamp- 
sil'is U(/a)nc'nfim(s, in many waters a poor or inditferent shell, is here 
remarkably large and thick, with nacre of an excellent luster, and is 
the best button species found in the river. This should be planted 
extensively as the main crop. Plagiola seci/ris, the butterfly, w^ould 
probably thrive in the water below Defiance Dam, and Sympliynota. 
costata would be well worth trying, especially in canals with black 
muddy bottoms, as it would be nearly certain to attain to unusual size 
and thickness there, and might rank as a good l)utton shell. 

The status of L. rectus is peculiar. In many streams and rivers this 
species is valueless on account of small size or purple nacre or both; 
in the Maumee it has large size and white nacre and is the most 
valuable species per pound native to the ri^er. With enough button 
shells in the river to bring this species to the market it would be de- 
sirable to plant rather heavily of it. It would make a good export 
shell and yield the clammer much more revenue for work done and 
material handled than the button species. 

The most valuable of the fresh-water mussels, Lampsilis anodon- 
toides, would be well worth a trial. It is not native to the river, but 
there is every reason to believe that, like its close relative L. rectus, 
it would grow and thrive considerably better even than in rivers 
where it is native. Good shells of this species bring an unusually 
high price on the market. 

As regards mussel parasites, the river appears to have about the 
average number as compared with other streams examined. They 
are usually most abundant in thin-shelled species in quiet waters. 
As regards pearls and pearly growths, wdiich are closely related to 
parasites, these are few. The distomid of Kelly and dorsal baroques 
are rare. Round pearls, all very small, were occasionally found, 
and all these appear to owe their origin to the marginal-cyst dis- 
tomid, which is quite common.** 

" Since this was written, we tiave found several minute pearls which were formed about 
eggs of Atax. 

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